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OF  THE 

UNIVERSITY 

Of    ILLINOIS 

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Ih.«,   citation  and  111  *>*»  •"'";' 


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Notes  on  Snakes  of  the  Genus  Calamaria 

Hymen  Marx 
Assistant,  Division  of  Reptiles  and  Amphibians 

AND 

Robert  F.  Inger 
Curator,  Division  of  Reptiles  and  Amphibians 

INTRODUCTION 

At  the  generic  level,  the  snakes  of  the  genus  Calamaria  are 
among  the  most  readily  recognized  of  the  entire  oriental  fauna. 
Yet  at  the  species  level,  they  present  a  bewildering  array  of  super- 
ficially similar  forms  that  have  been  for  the  most  part  poorly  defined. 
These  notes  are  offered  with  the  hope  that  our  analysis  of  the 
literature  and  discussion  of  characters  will  define  some  of  the  species 
and  suggest  fruitful  lines  of  attack  for  a  generic  revision. 

The  species  discussed  below  are  those  in  the  collections  of  Chicago 
Natural  History  Museum  (CNHM)  with  the  addition  of  Bornean 
forms  borrowed  from  other  institutions.  The  emphasis  on  Borneo 
arises  from  our  more  general  studies  of  the  Bornean  fauna.  We  are 
indebted  to  the  following  persons  for  the  loan  of  material  under 
their  care:  Mr.  CM.  Bogert,  American  Museum  of  Natural  History 
(AMNH);  Mr.  J.  C.  Battersby,  British  Museum  (Natural  History) 
(BM);  Mr.  Arthur  Loveridge,  Museum  of  Comparative  Zoology 
(MCZ);  Mr.  M.  W.  F.  Tweedie,  Raffles  Museum  (RM);  Miss 
Margaret  Storey,  Natural  History  Museum,  Stanford  University 
(SU);  and  Dr.  D.  M.  Cochran,  United  States  National  Museum 
(USNM).  We  are  grateful  to  Mr.  Harry  G.  Nelson,  Roosevelt 
University,  Chicago,  for  identification  of  some  food  fragments,  and  to 
Dr.  Rainer  Zangerl,  Chicago  Natural  History  Museum,  for  photo- 
graphic assistance.  The  drawings  are  the  work  of  Miss  Phyllis 
Wade,  Chicago  Natural  History  Museum. 

167 


168  FIELDIANA:  ZOOLOGY,  VOLUME  37 

SPECIALIZATIONS  OF  THE  GENUS  CALAMARIA 

According  to  published  accounts  (de  Rooij,  1917;  Taylor,  1922; 
Mertens,  1930;  Pope,  1935)  the  snakes  of  the  genus  Calamaria  are 
secretive,  generally  being  found  under  logs  and  other  debris  on  the 
ground. 

The  following  characters  define  the  external  morphological 
divergence  of  Calamaria  from  the  generalized,  freely  ranging  colubrid 
stock:  (1)  no  internasals;  (2)  prefrontals  broadly  in  contact  with 
supralabials;  (3)  reduction  in  size  of  nasal;  (4)  loreal  absent  (except 
in  C.  tropica  Taylor);  (5)  reduction  or  loss  of  oculars;  (6)  reduction 
in  size  of  eye;  (7)  reduction  in  number  of  labials;  (8)  parietals  in 
contact  with  supralabials;  (9)  reduction  in  overall  size;  (10)  reduction 
of  tail.  These  characters  add  up  to  consolidation  of  head  shields, 
proportional  reduction  of  head  and  tail,  and  reduction  of  overall 
size;  in  short,  the  modifications  commonly  associated  with  burrowing 
snakes.  Many  of  the  same  characters  are  found  in  Atractaspis, 
Atractus,  Typhlogeophis,  and  Sympholis. 

Different  levels  of  specialization  are  shown  by  the  various  species 
of  Calamaria.  But  all  forms  have  in  common  most  of  the  characters 
of  the  above  list.  Variation  in  the  degree  of  reduction  of  the  oculars 
and  eye,  the  number  of  labials,  the  length  of  the  snout,  and  the 
relative  proportions  of  head  shields  indicate  different  degrees  of 
specialization. 

Within  the  genus  the  diameter  of  the  eye  may  be  slightly  longer 
than,  equal  to  or  shorter  than  the  distance  of  the  eye  from  the 
mouth.  We  consider  that  the  smaller  the  eye  the  more  specialized 
the  form.  The  reduction  of  eye  size  is  associated  with  reduction 
in  the  preoculars.  We  have  tabulated  the  characters  of  80  currently 
recognized  species  (see  check  list)  of  Calamaria.  Data  were  taken 
from  the  literature;  the  original  descriptions  were  used  for  species 
not  included  in  the  major  faunal  works  (de  Rooij,  1917;  Taylor, 
1922;  Pope,  1935;  Smith,  1930,  1931,  1943).  The  relation  between 
the  presence  of  preoculars  and  the  size  of  the  eye  in  all  species  for 
which  data  are  available  is  given  in  Table  1.  It  is  clear  from  the 
table  that  the  disappearance  of  the  preocular  is  associated  with  the 
reduction  in  eye  size. 

Another  character  commonly  mentioned  in  the  literature  is  the 
ratio  of  the  width  of  the  frontal  to  the  width  of  the  supraocular. 
As  the  supraocular  becomes  smaller,  the  frontal  becomes  relatively 
wider.  This  ratio  is  also  associated  with  eye  size.  As  the  eye 
becomes  smaller  the  ratio  increases  (Table  2).    The  association  of 


MARX  AND  INGER:  SNAKES  OF  GENUS  CALAMARIA         169 

Table  1— ASSOCIATION  OF  PRESENCE  OF  PREOCULARS  WITH  SIZE 
OF  EYE  IN  SPECIES  OF  CALAMARIA 
Observed  numbers  of  species        Total         Calculated  numbers  of  species 


Preocular 

Preocular 

Present 

Absent 

Present 

Absent 

ye* 

Greater. 
Equal.  . 
Less 
Total . 

..   13 

.  .   23 

19 

..  55 

0 

4 

11 

15 

13 
27 
30 
70 

10.2 
21.2 
23.6 
55.0 

2.8 

5.8 

6.4 

15.0 

Chi  square=8.62;  n  =  2;  P=0.01 

*  "Greater,"  "equal,"  and  "less"  refer  to  eye  diameter  relative  to  distance 
of  eye  from  mouth. 

the  reduction  in  eye  size  with  the  reduction  of  the  preocular  and 
the  increase  of  the  frontal-supraocular  ratio  suggests  that  the  ocular 
region  represents  a  morphological  field  that  is  becoming  smaller 
as  a  whole. 

The  great  majority  of  species  of  Calamaria  have  five  or  six 
supralabials,  with  the  third  and  fourth  entering  the  eye.  About 
one-sixth  of  the  species  have  four  supralabials.  All  but  one  of  these 
have  the  second  and  third  supralabials  entering  the  eye.  Since 
most  colubrids  have  more  than  six  upper  labials,  those  Calamaria 
with  four  labials  are  considered  more  specialized  than  those  with 
five.  The  fact  that  the  change  from  five  to  four  is  apparent  in  front 
of  the  orbit  suggests  that  that  is  the  position  of  the  reduction. 
The  reduction  is  a  result  either  of  a  shortening  of  the  snout  or  of  a 
consolidation  of  the  shields  of  the  snout.  Each  of  these  processes 
seems  to  be  important  to  burrowing  snakes. 

The  reduction  in  the  number  of  labials  is  independent  of  eye 
size  (Table  3).  The  low  value  of  chi  square  in  Table  3  demonstrates 
that  these  two  characters  are  distributed  at  random,  relative  to  one 
another,  and  that,  therefore,  they  represent  different  morphological 
fields  with  different  selective  relationships. 

The  deviation  of  Calamaria  from  the  colubrid  pattern  of  scutel- 
lation  is  accompanied  by  modifications  of  the  skull  (fig.  21).  Some 
of  these  changes  parallel  the  consolidation  of  the  head  shields.  The 
discussion  that  follows  is  based  on  one  skull  each  of  C.  leucogaster, 
C.  gervaisi,  C.  leucocephala  and  C.  septentrionalis. 

In  most  non-specialized  colubrids  (as  in  Coluber,  Natrix,  Elaphe, 
and  Farancia)  the  septomaxilla  has  a  small  postero-lateral  projection 
that  is  widely  separated  from  both  prefrontal  and  nasal.  In  Cala- 
maria the  septomaxilla  has  a  lateral  wall,  subequal  in  length  to  the 


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MARX  AND  INGER:  SNAKES  OF  GENUS  CALAMARIA         171 

longitudinal  axis  of  the  nasal.  Posteriorly  this  broad  wall  abuts 
the  prefrontal  and  dorsally  has  an  extensive  connection  with  the 
lateral  border  of  the  nasal.  This  relationship  gives  the  septomaxilla 
the  position  of  a  strut  between  nasal  and  prefrontal  and,  presumably, 
adds  rigidity  to  the  snout.  This  rigidity  has  been  noticed  by  G. 
Haas  (1930,  p.  156),  who  comments  on  the  reduced  movability  of 
the  transverse  suture  between  the  nasals  and  the  frontals  in  Cala- 
maria  linnaei. 

The  walls  of  the  brain  case  are  also  modified.  The  parietal  and 
supraoccipital  above  and  the  basisphenoid  and  basioccipital  below 
lack  the  surface  sculpturing  of  most  colubrid  skulls.  Instead,  the 
brain  case  has  been  smoothed,  broadened,  and  flattened. 

But  far  more  radical  are  the  changes  that  have  taken  place  in 
some  of  the  bones  used  in  engulfing  food.  The  typical  snake  has  a 
long  squamosal  (about  equal  to  the  length  of  the  skull  behind  the 
frontal),  closely  applied  to  the  dorso-lateral  portion  of  the  parietal 
anteriorly  but  free  from  and  superior  to  the  occipital  region  pos- 
teriorly. Usually  (as  in  Coluber,  Elaphe,  Natrix,  and  Farancia)  the 
squamosal  projects  beyond  the  foramen  magnum.  The  quadrate  is 
as  long  as  or  longer  than  the  squamosal  and,  when  the  mouth  is 
closed,  slopes  backward  and  outward  from  its  articulation  with  the 
posterior  end  of  the  squamosal.  The  length,  position,  and  freedom 
of  movement  of  these  bones  are  necessary  to  the  great  mobility  of 
the  mandible. 

In  Calamaria  the  squamosal  is  reduced  to  a  tiny  sliver  no  longer 
than  the  dorsal  width  of  the  quadrate.  Instead  of  lying  on  the 
dorso-lateral  surface  of  the  parietal,  the  squamosal  lies  on  the  side 
of  the  skull  and  is  closely  applied  for  its  entire  length  to  the  prootic 
and  may  even  overlie  part  of  the  exoccipital.  The  quadrate,  closely 
pressed  against  the  side  of  the  skull  and  oriented  vertically,  is 
shorter  than  the  depth  of  the  brain  case;  its  dorso-ventral  axis  is 
subequal  to  its  dorsal  width.  The  posterior  edge  of  the  quadrate 
lies  anterior  to  the  level  of  the  foramen  magnum.  The  shortening 
and  the  changed  positions  of  the  squamosal  and  the  quadrate  in 
Calamaria  must  reduce  the  amplitude  of  jaw  movement. 

The  pterygoid  of  Calamaria  is  also  shortened,  probably  in  asso- 
ciation with  the  changed  position  of  the  quadrate.  Whereas  the 
pterygoid  in  colubrids  usually  projects  far  beyond  the  level  of  the 
foramen  magnum,  in  Calamaria  it  ends  at  that  level  or  anteriorly. 

The  shortening  of  these  three  bones  may  be  viewed  as  part  of 
the  process  of  consolidation.    With  the  confinement  of  the  ends  of 


Fig.  21.    Skull  of  Calamaria  leucogaster  (CNHM  71598);  actual  length  7  mm. 


172 


MARX  AND  INGER:  SNAKES  OF  GENUS  CALAMARIA 


173 


the  pterygoid  and  squamosal  to  the  limits  of  the  brain  case  and  the 
pulling  in  of  the  quadrate  against  the  skull,  some  of  the  "loose 
ends"  of  the  skull  have  been  eliminated  and  the  three  bones  have 
been  given  added  protection. 


Fig.  22.    Maxilla  of  Calamaria  leucogaster  (CNHM  71598) ;  actual  length  5  mm. 
First  tooth  missing. 


One  additional  specialization  of  Calamaria  is  unique.  The 
maxillary  teeth  of  all  but  two  of  the  species  examined  are  greatly 
broadened  at  the  base  and  compressed  laterally  (figs.  22-24).  Near 
the  apex  of  the  enlarged  base  is  a  small,  oval,  translucent  area.  The 
pulp  cavity  (fig.  24)  has  two  branches,  one  running  out  into  the 


Fig.  23.    Maxilla  of  Calamaria  septentrionalis  (CNHM  7139);  actual  length 
3.5  mm.    Last  two  teeth  missing. 


shaft  and  the  other  curving  into  the  posterior  part  of  the  base. 
From  the  saddle  between  these  two  branches,  fine  pulp  canals  run 
into  the  oval,  translucent  "window."  The  small  amount  of  pulp  in 
this  region  is  apparently  responsible  for  its  transparency. 


174  FIELDIANA:  ZOOLOGY,  VOLUME  37 

Two  levels  of  maxillary  tooth  specialization  are  found  in  the 
species  we  have  examined.  The  less  modified  condition  occurs  in 
schmidti,  which  has  seven,  widely  spaced,  unmodified  ophidian  teeth, 
and  in  leucogaster,  in  which  only  the  last  two  of  its  seven  maxillary 
teeth  are  slightly  enlarged  at  the  base  (fig.  22).  The  more  specialized 
condition  is  characteristic  of  all  the  others.  In  these  last,  all  of  the 
maxillary  teeth  are  modified  as  illustrated.  There  is  no  space 
between  successive  teeth,  and  all  are  enlarged  at  the  base. 

The  other  dentigerous  bones  have  typical  ophidian  teeth  (fig.  25). 


Fig.  24.  Maxillary  tooth  of  Calamaria  gervaisi  (CNHM  15025),  greatly  en- 
larged; cleared  and  stained  with  eosin. 

The  function  of  the  specialized  teeth  is  obscure.  The  diet  of 
Calamaria  consists  largely,  if  not  exclusively,  of  earthworms.  Four- 
teen individuals,  belonging  to  eight  of  the  eighteen  species  studied, 
contained  earthworms.  There  were  no  marks  on  the  food  remnants 
to  indicate  how  the  teeth  were  used.  Other  earthworm  feeders 
among  oriental  colubrids — Natrix  nuchalis,  Achalinus  spinalis, 
Opheodrys  major,  and  Trirhinopholis  styani  (Pope,  1935) — have  the 
simple,  conical,  rather  widely  spaced  teeth  characteristic  of  snakes — 
an  indication  that  the  unusual  teeth  of  Calamaria  may  not  be  an 
adaptation  to  food  itself. 

Instead,  G.  Haas'  description  (1930,  p.  157)  of  the  skull  move- 
ments of  C.  linnaei  causes  us  to  speculate  that  the  modified  teeth 
may  constitute  an  adaptation  to  facilitate  food  ingestion.  Though 
not  commenting  on  the  form  of  the  maxillary  teeth,  Haas  concludes 
that  the  maxilla  is  capable  only  of  very  limited  movement  in  a 
longitudinal  direction.    Rather,  during  protraction  of  the  pterygo- 


MARX  AND  INGER:  SNAKES  OF  GENUS  CALAMARIA         175 

palatal  arch,  the  maxilla  is  swung  laterally  so  that  the  tooth  row, 
which  at  rest  is  oriented  obliquely  inward,  is  rotated  on  its  longi- 
tudinal axis.  Haas  suspects  that  this  swinging  action  of  the  maxilla 
functions  to  push  the  prey,  whose  soft  body  tends  to  be  squeezed 


Fig.  25.    Mandible  of  Calamaria  septentrionalis  (CNHM  7139) ;  actual  length 
8.5  mm. 


out  laterally  by  the  pressure  of  jaw  and  palate,  in  toward  the  center 
of  the  mouth  so  that  it  can  pass  the  gape. 

Assuming  Haas  to  be  correct,  it  seems  clear  that  the  broadening  of 
the  base  of  each  maxillary  tooth  would  increase  the  effectiveness 
of  the  maxilla  by  increasing  the  surface  it  could  bring  to  bear  on 
the  prey.  Furthermore,  since  Haas  states  that  the  turning  of  the 
maxilla  is  stronger  posteriorly,  we  are  presented  with  a  functional 
explanation  for  the  observation  that  apparently  the  posterior 
maxillary  teeth  were  the  first  to  become  broadened  (as  in  leuco- 
gaster). 

EVALUATION  OF  CHARACTERS 

Since  80  species  of  Calamaria  are  now  recognized,  part  of  the 
difficulty  of  identifying  these  snakes  lies  in  the  number  of  species 
to  be  compared.  Unfortunately,  some  authors  give  the  impression 
of  extreme  haste  in  describing  new  forms;  it  almost  seems  that  the 
mass  of  forms  to  be  compared  discouraged  any  comparison.  But 
most  of  the  trouble  arises  from  inadequate  definition  of  the  species 
in  the  literature.  In  general,  the  literature  fails  to  evaluate  the 
characters.  The  significance  of  a  range  of  fifty  ventrals,  for  example, 
is  not  examined.  Indeed,  many  authors  have  neglected  to  sex  their 
snakes,  thus  rendering  their  counts  even  less  useful. 

The  characters  most  often  used  to  distinguish  the  species  are: 
(1)  the  number  of  supralabials;  (2)  the  contact  or  separation  of  the 
mental  and  first  chin  shields;  (3)  the  ratio  of  frontal  width  to  supra- 


176  FIELDIANA:  ZOOLOGY,  VOLUME  37 

ocular  width;  (4)  the  diameter  of  the  eye  relative  to  its  distance  from 
the  mouth;  (5)  the  presence  or  absence  of  a  preocular;  and  (6)  the 
number  of  ventrals  (see  Boulenger,  1894,  and  de  Rooij,  1917). 

The  number  of  supralabials  varies  from  four  to  six.  The  dif- 
ference between  five  and  six  is  merely  a  difference  in  the  method  of 
counting  and  depends  upon  whether  the  small  scale  following  the 
large  fifth  labial  is  included  in  the  count;  in  either  case  the  same 
labials  enter  the  eye  and  the  same  labial  (fifth)  is  the  largest.  No 
significance  can  be  attached  to  the  difference  between  five  and  six 
labials  and  no  forms  are  so  distinguished.  The  distinction  between 
four  and  five  labials,  however,  is  important.  All  forms  with  five 
(or  six)  labials  have  the  third  and  fourth  scutes  entering  the  orbit, 
whereas  all  those  with  four  labials  have  the  second  and  third  enter- 
ing the  orbit,  with  the  exception  of  lowi  in  which  only  the  third 
enters  it. 

We  have  examined  two  or  more  specimens  of  each  of  fifteen 
species;  the  two  largest  samples  were  47  (vermiformis)  and  20  (ger- 
vaisi).  No  intra-specific  variation  in  supralabial  counts  was  ob- 
served, nor  is  there  any  mention  of  such  variation  in  the  literature. 
A  form  invariably  has  either  four  or  five  (or  five  or  six)  supralabials. 

Almost  as  constant  is  the  contact  or  separation  of  the  mental 
and  anterior  chin  shields.  We  have  seen  only  one  instance  of  intra- 
specific  variation.  Normally  the  first  infralabials  meet  behind  the 
mental  in  C.  leucogaster,  but  one  of  the  four  examined  has  the  mental 
narrowly  in  contact  with  the  chin  shields.  Thus,  this  character  on 
the  whole  is  reliable  in  distinguishing  species,  but  caution  must  be 
used  when  only  a  single  specimen  is  available. 

The  ratio  of  frontal  width  to  supraocular  is  subject  both  to  in- 
dividual variation  and  to  errors  in  measurement.  The  ratio  in  the 
large  series  of  vermiformis  varied  from  \Yi  to  23^;  gervaisi  varied 
from  13^  to  2;  four  leucocephala  showed  a  range  of  4  to  5.  Clearly, 
a  difference  of  1  in  this  ratio  can  easily  be  accounted  for  by  indi- 
vidual variation,  but  a  difference  of  2  or  more,  as  in  the  comparison 
of  gervaisi  and  leucocephala,  is  a  matter  of  species  differentiation. 
Keys  (for  example,  Boulenger,  1894;  de  Rooij,  1917)  that  separate 
groups  of  species  on  the  basis  of  the  frontal  being  twice  or  less  than 
twice  as  wide  as  the  supraocular  are  not  very  helpful  except  in  the 
extremes. 

Much  the  same  can  be  said  for  eye  size.  There  are  species  with 
extremely  small  eyes  and  those  with  moderately  large  eyes.  Some 
intra-specific  variation  is  evident  but  it  never  includes  the  extremes 


MARX  AND  INGER:  SNAKES  OF  GENUS  CALAMARIA         177 

of  the  genus.  In  general,  only  when  species  occupy  different  ends 
of  this  scale  can  they  be  distinguished  on  the  basis  of  this  character. 
The  presence  or  absence  of  a  preocular  is  relatively  constant. 
The  notable  exception  in  our  experience  is  the  series  of  leucocephala. 
One  of  the  four  seen  has  no  preocular  on  either  side  of  the  head;  a 
second  has  a  preocular  on  one  side  only;  the  other  two  have  a  pre- 
ocular on  both  sides.  But  when  present  the  preocular  is  very  small 
and  the  prefrontal  has  a  broad  contact  with  the  eye.  Apparently 
the  preocular  is  in  the  process  of  disappearing  in  leucocephala.  In 
species  with  moderately  large  preoculars,  the  absence  of  this  scale 
is  probably  a  rare  anomaly. 

The  range  of  ventral  counts  reported  in  the  literature  is  presented 
uncritically  for  the  most  part.  Even  Boulenger  (1894),  one  of  the 
few  authors  to  sex  his  specimens,  failed  to  analyze  the  counts;  for 
example,  he  gave  153-210  as  the  range  of  ventral  counts  for  vermi- 
formis  but  did  not  comment  on  this  range  or  determine  it  for  a  single 
series.  Analysis  of  our  sample  of  vermiformis  (snakes  from  the  Malay 
Peninsula,  Sumatra,  Borneo,  and  the  Philippine  Islands)  provides 
an  estimate  of  what  is  probably  near  the  maximum  intra-specific 
variation  for  the  genus.  The  total  range  of  ventrals  for  our  22  males 
is  53  (144-196)  and  for  the  25  females  81  (137-217).  Even  if  only 
Bornean  snakes  are  considered,  the  range  for  12  females  is  64 
(137-200).  It  is  only  when  analysis  is  limited  to  a  single  locality 
that  the  variation  is  not  unusually  great;  for  example,  our  Bornean 
sample  includes  five  males  and  five  females  from  Mount  Kina  Balu; 
their  ventral  counts  are,  respectively,  154-165  and  170-180,  ranges 
that  are  very  narrow  in  contrast  to  the  total  range  for  the  species. 
Similarly,  although  the  four  Sumatran  males  examined  have  a  ventral 
range  of  31  (166-196),  three  from  a  single  locality  vary  only  from 
191  to  196;  seven  Sumatran  females  from  scattered  areas  have  a 
ventral  range  of  39  (179-217),  whereas  four  from  one  locality  have 
a  range  of  only  9  (209-217). 

This  pattern  of  wide  intra-specific  variation  coupled  with  narrow 
local  variation  is  also  found  in  other  species.  In  gervaisi,  the  total 
ventral  range  of  22  females  from  Luzon,  Negros,  and  Mindanao  is 
39  (142-180);  but  nine  from  Montalban,  Luzon,  show  a  range  of 
only  9  (158-166).  Five  males  of  borneensis  vary  from  152  to  169, 
yet  four  from  one  locality  have  a  range  of  only  11  (159-169). 

Given  such  wide  intra-specific  variation,  ventral  counts  are  of 
little  use  in  distinguishing  forms.  It  is  only  when  allopatric  popu- 
lations differ  by  more  than  50  ventrals  that  any  significance  can  be 


178 


FIELDIANA:  ZOOLOGY,  VOLUME  37 


attached  to  the  difference.  On  the  other  hand,  in  dealing  with 
closely  related  sympatric  species,  a  difference  of  20  ventrals  can  be 
accepted  as  an  inter-specific  difference  if  both  samples  come  from 
the  same  restricted  area  and  if  the  sexes  are  handled  separately. 

Although  the  absolute  variation  in  subcaudal  counts  must  be 
smaller  because  of  the  numbers  involved,  relatively  they  show  the 
same  range  as  the  ventrals.  The  range  of  81  ventrals  in  vermiformis 
equals  37  per  cent  of  the  maximum  count  (217).  The  subcaudals 
of  our  25  female  vermiformis  vary  from  14  to  20;  this  range  of  7  is 
equal  to  35  per  cent  of  the  highest  count.  Yet  despite  this  wide 
intra-specific  variation,  there  is  much  inter-specific  differentiation 
in  caudal  counts  (see  Table  4).  However,  because  of  a  very  sharp 
sex  dimorphism,  subcaudal  counts  are  worthless  unless  the  sex  is 
determined. 

Table  4.— POSITION  OF  REDUCTION  TO  FOUR  DORSAL  SCALE 
ROWS  IN  CERTAIN  SPECIES  OF  CALAMARIA 

Number  of  specimens  given  in  parentheses 

Males  Females 

Total  Total 

Species  Position*  subcaudals  Position  subcaudals 

leucogaster 4-8  (4)  16-18  (4) 

gervaisi 6-11  (3)  16-19  (6)  2-10  (16)  11-16  (22) 

griswoldi 9-12  (4)  16-17  (4)  9-10  (3)  14-16  (3) 

lumbricoidea...      4-5(2)  19-22(2)                  5(1)                       17(1) 

vermiformis.  .  .      2-9  (22)  19-27  (22)  0-7  (25)  14-20  (25) 

grabowskyi.  .  .  .     16-19  (2)  26-27  (2)  14-18  (2)  22-23  (2) 

suluensis 9(2)  23-25(2) 

pendleburyi . . . .       9(1)  18(1)  12(1)                      17(1) 

lautensis 8  (1)  19  (1)                     6  (1)                       12  (1) 

leucocephala . .  .     12-15(2)  34-37(2)  8-12(2)  26-29(2) 

septentrionalis.        0  (1)  18  (1)                     0  (2)  10-11  (2) 

borneensis 6-9  (4)  22-24  (5)                  4  (3)  16-19  (3) 

schmidti 6-7  (2)  14  (2) 

loivi 4-6  (2)  19-20  (2)  3-5  (2)                    14  (2) 

*Position  located  in  terms  of  subcaudals,  counting  forward  from  terminal 
scute. 


Another  character,  not  discussed  in  the  literature,  has  been 
useful  in  distinguishing  certain  species.  The  number  of  dorsal  scale 
rows  on  the  tail  is  usually  reduced  to  four  shortly  before  the  terminal 
scute.  The  point  at  which  the  reduction  to  four  scale  rows  occurs, 
as  located  by  the  number  of  subcaudals  counted  forward  from  the 
terminal  scute,  is  relatively  constant  within  species  (Table  4).  The 
position  of  this  reduction  seems  to  depend  more  on  the  shape  of  the 


MARX  AND  INGER:  SNAKES  OF  GENUS  CALAMARIA         179 

tail  than  on  its  length,  as  is  shown  in  Table  4  by  a  comparison  of 
grabowskyi  with  vermiformis  and  of  lautensis  with  septentrionalis. 
The  data  of  Table  4  demonstrate  that  it  would  be  difficult  to  identify 
a  single  specimen  on  the  basis  of  this  character  alone,  but  it  is  useful 
in  substantiating  conclusions  arrived  at  on  the  basis  of  other  charac- 
ters. We  refer  to  the  comparisons  of  griswoldi  with  vermiformis 
grayi  (see  p.  184)  and  of  grabowskyi  with  suluensis  (see  p.  191). 

TAXONOMIC  SECTION 

Calamaria  leucogaster  Bleeker.    Figures  21,  22. 

Calamaria  leucogaster  Bleeker,  1860,  Nat.  Tijds.  Ned.  Indie,  21 :  293 — Ampat- 

Lawang,  Sumatra. 
Calamaria  brookii  Boulenger,  1895,  Ann.  Mag.  Nat.  Hist.,  (6),  15:  329. 
Calamaria  brookei  de  Rooij,  1917,  Rept.  Indo-Austr.  Arch.,  2:  166. 

Descriptive  notes. — Four  females;  total  length  178-208  mm.;  tail 
0.06-0.08  of  total;  ventrals  146-155;  subcaudals  16-18;  supralabials 
5,  third  and  fourth  entering  eye;  infralabials  5,  first  three  pairs  in 
contact  with  anterior  chin  shields;  oculars  1+1;  position  of  reduction 
to  4  dorsal  scale  rows,  counting  subcaudals  forward  from  terminal 
scute  4-8;  maxilla  with  7  conical  teeth. 

Discussion. — One  (CNHM  67279)  differs  from  the  others  in 
having  a  narrower  snout  and  in  having  the  mental  in  contact  with 
the  chin  shields  instead  of  being  separated  by  the  first  infralabials. 
The  latter  distinction  has  been  thought  significant  in  distinguishing 
species  of  Calamaria.  This  is  the  only  example  of  individual  varia- 
tion in  this  character  that  we  have  found.  The  first  infralabials  of 
CNHM  67279  are  only  narrowly  separated.  Otherwise  all  four 
snakes  are  so  similar  that  we  must  place  them  in  the  same  species. 
Especially  significant  is  the  agreement  in  maxillary  dentition  (see 
discussion,  p.  174). 

The  differences  between  leucogaster  and  brooki  as  determined 
from  Boulenger's  descriptions  (1894,  1895),  which  were  based  upon 
examination  of  the  types,  are:  (1)  frontal  of  leucogaster  twice  as 
wide  as  supraocular,  three  times  as  wide  as  supraocular  in  brooki; 
(2)  snout  of  leucogaster  pointed,  no  description  for  brooki;  (3)  sub- 
caudals of  leucogaster  14-17,  of  brooki  23. 

As  noted  above,  one  of  our  snakes  (CNHM  67279)  has  a  more 
pointed  snout  than  the  others.  In  this  character,  therefore,  it 
resembles  leucogaster.  But  the  frontal  of  this  specimen  relative  to 
the  supraocular,  said  to  be  wider  in  brooki,  is  wider  than  the  others. 


180  FIELDIANA:  ZOOLOGY,  VOLUME  37 

Accordingly,  our  material  shows  a  recombination  of  these  supposedly- 
diagnostic  characters.  The  differences  in  subcaudals  can  be  at- 
tributed to  sex  dimorphism;  the  type  of  brooki  is  a  male,  whereas 
Boulenger  (1894)  gives  no  counts  for  male  leucogaster. 

In  Boulenger's  descriptions,  these  two  forms  also  differ  in 
coloration.  The  difference  lies  in  the  apparent  absence  of  longitu- 
dinal dark  lines  in  leucogaster.  But  de  Rooij  (1917),  who  also 
examined  the  type  of  leucogaster,  notes  that  eight  dark  longitudinal 
lines  are  present.  Boulenger  mentions  only  five  in  brooki  plus  a 
dark  upper  edge  on  the  outer  scale  rows.  If  that  dark  edge  be 
counted  as  an  additional  line,  the  total  in  brooki  is  raised  to  seven. 
The  median  band  in  our  material  is  split  into  two  by  a  light  mid- 
dorsal  line.  Otherwise  the  agreement  between  our  snakes  and  Bou- 
lenger's description  of  brooki  is  so  close  that  we  assume  we  are  dealing 
with  the  same  form,  and,  therefore,  that  the  differences  in  the  mid- 
dorsal  stripe  can  be  explained  as  individual  variation. 

Since  none  of  the  distinctions  between  leucogaster  and  brooki 
stand  up  under  analysis  we  must  place  the  latter  in  synonymy. 
This  conclusion  is  supported  by  the  collection  of  both  nominate 
forms  from  Matang,  Sarawak  (de  Rooij,  1917,  pp.  164,  166). 

Localities  and  specimens  examined. — North  Borneo:  Jesselton 
District:  Mount  Kina  Balu,  Kiau,  3,000  feet  (RM  320). 

Sarawak:  First  Division:  Santubong,  Sungai  Ja'ong  (CNHM 
71598-99).    Fifth  Division:  Lawas  (CNHM  67279). 

Calamaria  gervaisi  DumeYil  and  Bibron.    Figure  24. 

Calamaria  virgulata  (non  Boie)  Eydoux  and  Gervais,  1837,  in  Guer.  Mag. 
Zool.,  1837,  CI.  3,  pi.  16,  figs.  7-10. 

Calamaria  gervaisi  Dumeril  and  Bibron,  1854,  Erp.  Gen.,  7:  76 — Java  [in 
error,  see  remarks,  p.  182]. 

Descriptive  notes. — Twenty  specimens;  supralabials  5,  third  and 
fourth  entering  eye;  infralabials  5  or  6,  first  pair  not  in  contact 
behind  mental,  first  3  pairs  (18),  4  pairs  (1),  or  %  pairs  (1)  in  contact 
with  anterior  chin  shields;  oculars  1  +  1;  maxilla  with  8  or  9  modified 
teeth;  for  ventral  and  subcaudal  counts  see  Table  5. 

Luzon:  Two  males;  total  length  190-207  mm.;  tail  0.07-0.08 
of  total.  Nine  females;  total  length  140-269  mm.;  tail  0.04-0.05  of 
total.  Position  of  reduction  to  4  dorsal  scale  rows,  counting  sub- 
caudals forward  from  terminal  scute  2-7  (mean  5.1). 


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182  FIELDIANA:  ZOOLOGY,  VOLUME  37 

Negros:  Two  females;  total  length  244-296  mm.;  tail  0.05-0.06 
of  total.  Position  of  reduction  to  4  dorsal  scale  rows,  counting 
subcaudals  forward  from  terminal  scute  5-6. 

Mindanao:  One  male;  total  length  179  mm.;  tail  0.09  of  total. 
Five  females;  total  length  158-258  mm.;  tail  0.05-0.07  of  total. 
One  unsexed.  Position  of  reduction  to  4  dorsal  scale  rows,  counting 
subcaudals  forward  from  terminal  scute  7-11  (mean  9.1). 

Discussion. — On  the  basis  of  a  greater  number  of  ventrals, 
Taylor  (1917)  distinguished  the  Negros  population  as  the  subspecies 
g.  iridescens.  Our  material  bears  out  the  difference  in  this  character. 
The  Mindanao  population  apparently  differs  from  both  the  Luzon 
and  the  Negros  populations  in  the  same  character  (see  Table  5). 
Furthermore,  the  Mindanao  animals  have  a  yellow  collar,  which  is 
not  found  in  any  of  our  Luzon  or  Negros  snakes  and  is  not  mentioned 
by  either  Boulenger  (1894,  p.  338)  or  Taylor  (1922,  p.  186).  The 
Mindanao  series  also  differs  from  the  others  in  the  position  of  the 
final  scale  row  reduction  (see  above) . 

If  the  Negros  population  is  recognized  as  a  distinct  subspecies, 
probably  the  Mindanao  population  should  be  also.  But  a  decision 
on  the  definition  of  subspecies  must  wait  for  additional  material 
from  scattered  localities. 

The  type  locality  of  gervaisi  is  obscure.  Dumeril  and  Bibron 
(1854)  state  that  Eydoux  and  Gervais  collected  their  snake,  the 
type  of  gervaisi,  in  Java.  Eydoux  and  Gervais  (1837)  give  no 
locality,  but  the  other  reptiles  that  they  specifically  state  having 
collected  are  all  from  Manila.  The  figure  of  Jan  and  Sordelli  (1865) 
of  the  type  shows  a  Calamaria  without  a  light  collar  and  with  a 
uniform  venter.  This  combination  is  characteristic  of  the  Luzon 
population,  so  the  type  probably  came  from  Luzon.  All  subsequent 
authors  (Gunther,  1858;  Boulenger,  1894;  Taylor,  1922)  restrict 
gervaisi  to  the  Philippines.  De  Haas  (1950)  does  not  include  it  in 
the  fauna  of  Java. 

Earthworms  were  found  in  the  guts  of  three  snakes  from  Luzon 
and  two  from  Mindanao. 

Localities  and  specimens  examined. — Philippine  Islands:  Luzon: 
Manila  (CNHM  15025-31,  15033-35);  Montalban  (CNHM  15036). 
Mindanao:  Mount  McKinley,  3,000  feet  (CNHM  53374-76) ;  Mount 
Apo,  2,800  feet  (CNHM  53377);  Tagum,  sea  level  (CNHM  53378- 
80).  Negros:  Dumaguete  (CNHM  57308);  Bais,  Mabaha  (CNHM 
61625). 


MARX  AND  INGER:  SNAKES  OF  GENUS  CALAMARIA         183 

Calamaria  lumbricoidea  Boie 

Calamaria  lumbricoidea  Boie,  1827,  Isis,  20:  540 — Java. 

Descriptive  notes. — Two  males;  total  length  428-468  mm.;  tail 
0.06-0.07  of  total;  ventrals  186-190;  subcaudals  19-22.  One  female; 
total  length  505  mm.;  tail  0.05  of  total;  ventrals  192;  subcaudals  17. 

Supralabials  5,  third  and  fourth  in  contact  with  eye;  infralabials 
5,  first  three  pairs  in  contact  with  anterior  chin  shields,  first  pair 
not  in  contact  behind  mental;  oculars  1  +  1;  position  of  reduction  to 
4  dorsal  scale  rows,  counting  subcaudals  forward  from  terminal 
scute  4-5;  maxilla  with  10-11  modified  teeth. 

In  coloration  these  agree  with  Boulenger's  description  (1894). 

Reasons  for  separating  griswoldi  from  lumbricoidea  are  given 
below. 

Localities  and  specimens  examined. — Java  :  Benkeng,  near  Serang 
(AMNH  71513).    No  other  data  (MCZ  25894). 

Sumatra:  No  other  data  (USNM  56439). 

Calamaria  griswoldi  Loveridge 

Calamaria  lumbricoidea  griswoldi  Loveridge,  1938,  Proc.  Biol.  Soc.  Washington, 
51:  43 — Luidan  River,  Bundu  Tuhan,  Mount  Kina  Balu,  North  Borneo. 

Descriptive  notes. — Four  males;  total  length  236-425  mm.;  tail 
0.06  (4)  of  total;  ventrals  166-179;  subcaudals  16-17.  Three  females; 
total  length  375-467  mm.;  tail  0.05-0.06  of  total;  ventrals  183-190; 
subcaudals  14-16. 

Supralabials  5,  third  and  fourth  entering  eye;  infralabials  5,  first 
pair  not  in  contact  behind  mental,  first  three  pairs  in  contact  with 
anterior  chin  shields;  oculars  1+1;  position  of  reduction  to  4  dorsal 
scale  rows,  counting  subcaudals  forward  from  terminal  scute  9-12; 
maxilla  with  10  modified  teeth. 

Discussion. — These  animals,  two  of  which  are  topotypes,  agree 
in  all  significant  details  with  Loveridge's  description. 

There  is  little  difficulty  in  separating  griswoldi  from  vermiformis, 
lumbricoidea,  or  grayi.  Superficially  the  dorsal  coloration  of  gris- 
woldi is  much  like  that  of  grayi  and  the  similarly  striped  "high 
altitude"  form  of  vermiformis,  but  in  grayi  and  vermiformis  the  light 
area  covers  only  the  edges  of  the  mid-dorsal  scales,  forming  very 
narrow  light  lines,  whereas  in  griswoldi  the  lateral  fourths  of  each 
scale  are  light,  forming  mid-dorsal  stripes  equal  to  half  the  width  of 
the  dark  interspaces.  The  immaculate  yellow  or  white  venter  of 
griswoldi  is  radically  different  from  the  boldly  marked  one  of  grayi 


184  FIELDIANA:  ZOOLOGY,  VOLUME  37 

and  vermiformis.  The  ventral  coloration  of  griswoldi  is  like  that  of 
lumbricoidea,  but  the  latter  is  uniformly  dark  mid-dorsally,  with  a 
yellow  stripe  running  the  length  of  the  outer  one  or  two  scale  rows. 

The  eye  of  griswoldi  is  distinctly  larger  than  the  eyes  of  the 
other  forms,  its  diameter  being  at  least  equal  to  its  distance  from  the 
mouth  but  usually  larger.  The  eye  diameter  is  usually  smaller  than 
its  distance  from  the  mouth  in  the  other  forms,  but  may  be  equal  to 
that  distance. 

The  reduction  to  4  dorsal  scale  rows  occurs  opposite  the  ninth 
to  twelfth  subcaudal  from  the  tip  of  the  tail  in  griswoldi  (mean  of 
7  snakes=9.7).  In  vermiformis  the  range  of  this  point  is  0-9  sub- 
caudals  from  the  tip  (only  one  with  more  than  7;  mean  of  39=4.3), 
in  grayi  3-6  (mean  of  8=5.0),  and  in  lumbricoidea  4-5  (mean  of 
3=4.7). 

Conceivably,  if  the  total  number  of  subcaudals  were  decreased, 
the  distance  of  the  final  reduction  from  the  tip  of  the  tail  in  terms 
of  subcaudals  should  become  less  and  less,  all  other  things  being 
equal.  But  the  difference  between  griswoldi  and  the  others  in  this 
distance  is  independent  of  total  subcaudal  count  as  griswoldi  has 
fewer  subcaudals. 

Subcaudal  counts  of  griswoldi  are  lower  than  those  of  the  other 
three  forms.  Observed  ranges  for  males  are:  griswoldi  16-17  (four); 
vermiformis  19-26  (seventeen),  only  one  with  less  than  22;  lumbri- 
coidea 19-22  (two);  grayi  22-27  (four).  Observed  ranges  for  females 
are:  griswoldi  13-16  (four,  including  the  type);  vermiformis  14-20 
(twenty-one),  only  four  with  less  than  17;  lumbricoidea  17  (one); 
grayi  15-18  (four),  only  one  under  17. 

Ten  vermiformis  from  the  type  locality  of  griswoldi  (Mount  Kina 
Balu)  differ  from  the  latter  in  ventral  counts.  The  five  males  of 
this  series  of  vermiformis  have  ventral  counts  of  154-165  and  the 
females  counts  of  170-180.  Two  male  griswoldi  have  170  and  179 
ventrals  and  five  females  (including  the  type  series)  a  range  of 
183-192.  This  difference  between  the  two  species  does  not  hold  up 
in  other  parts  of  the  geographic  range  of  vermiformis,  which  has  a 
total  spread  of  144-196  in  the  males  and  137-217  in  the  females. 

Though  originally  described  as  a  subspecies  of  lumbricoidea, 
griswoldi  differs  from  the  latter  sufficiently  in  eye  size,  position  of 
caudal  scale  reduction,  scale  counts,  and  coloration  to  warrant 
recognition  as  a  distinct  species. 

The  male  from  Lumu  Lumu,  Mount  Kina  Balu,  identified  by 
Smith  (1931)  as  vermiformis,  is  griswoldi,  as  is  clear  from  its  colora- 
tion and  counts. 


MARX  AND  INGER:  SNAKES  OF  GENUS  CALAMARIA         185 

One  snake  contained  an  entire  earthworm  (78  mm.  total  length) 
of  the  subfamily  Megascolecinae.  The  gut  of  the  earthworm  was 
full  of  wood  fragments  but  held  no  mineral  particles.  Apparently, 
the  earthworm  was  feeding  in  a  decaying  log  and  was  caught  there 
by  the  snake. 

Localities  and  specimens  examined. — North  Borneo:  Jesselton 
District:  Mount  Kina  Balu,  Bundu  Tuhan,  4,500  feet  (USNM 
130238-39);  Mount  Kina  Balu,  Lumu  Lumu,  5,000  feet  (BM 
1929.12.22.112);  Mount  Kina  Balu,  Tenompok,  4,700  feet  (RM 
18949,  RM  no  number);  Mount  Kina  Balu  (BM  95.11.7-25); 
Ranau  (USNM  134114). 

Calamaria  vermiformis  Dumeril  and  Bibron 

Discussion. — We  place  bruegeli  in  the  synonymy  of  vermiformis 
because  the  differences  noted  by  Mertens  are  within  the  range  of 
individual  variation.  According  to  Mertens,  the  frontal  of  bruegeli 
is  twice  as  wide  as  the  supraocular;  Boulenger  (1894)  states  that 
the  frontal  of  vermiformis  is  not  twice  as  broad  as  the  supraocular. 
Our  own  specimens  span  this  minor  difference.  The  other  distinction 
noted  by  Mertens  is  the  relatively  broader  rostral  of  bruegeli.  Due 
to  the  differences  of  methods  of  measuring  and  individual  variation 
in  our  specimens  this  distinction  does  not  warrant  recognition  of  a 
new  species.  Furthermore,  the  counts  and  other  characters  of 
bruegeli  agree  with  the  original  description  of  vermiformis  and  with 
our  material. 

The  Philippine  form,  grayi,  is  obviously  closely  related  to  vermi- 
formis in  counts,  habitus,  and  scale  proportions.  The  ventral 
coloration  of  grayi — barred  with  black — is  just  like  that  of  Bornean 
vermiformis.  Most  striking,  however,  is  the  agreement  in  juvenile 
pattern.  Two  young  grayi,  measuring  149  mm.  (MCZ  25781)  and 
216  mm.  (MCZ  25778),  have  the  yellow  heads  and  yellow  rings 
typical  of  juvenile  vermiformis,  but  the  lined  pattern  of  adult  grayi 
is  clearly  visible  in  the  dark  interspaces. 

Minor  differences  do  distinguish  the  two  populations.  Most 
outstanding,  of  course,  is  the  dorsal  coloration,  consisting  in  adult 
grayi  of  narrow  light  lines  on  a  dark  background  and  in  adult  Bornean 
vermiformis  of  a  uniform,  brown  color.  In  other  populations  of 
vermiformis,  for  example,  from  the  Malay  Peninsula  and  Sumatra, 
occasional  individuals  have  the  "grayi"  pattern. 

The  Philippine  form  seems  to  have  higher  ventral  counts  than 
the  Bornean  and  Malayan  populations  but  agrees  with  Sumatran 
vermiformis  (see  data  below). 


186  FIELDIANA:  ZOOLOGY,  VOLUME  37 

Excluding  the  last  labial,  each  parietal  of  grayi  is  usually  bordered 
postero-laterally  by  4  scales;  in  only  one  of  the  eight  seen  was  each 
parietal  bordered  by  3  scales.  On  the  other  hand,  only  one  out  of 
23  Bornean  vermiformis  and  one  out  of  11  Sumatran  vermiformis 
have  4  scales  in  contact  with  each  parietal;  all  the  others  have  3 
scales  bordering  each  parietal. 

In  view  of  the  great  similarity  between  grayi  and  vermiformis, 
we  must  consider  them  conspecific.  However,  the  differences  cited 
above  between  grayi  and  Bornean  vermiformis  justify  the  separation 
of  these  as  distinct  subspecies. 

The  remaining  samples  of  vermiformis  we  have  seen  show  a 
complex  pattern  of  geographic  variation.  Some  characters,  such 
as  ventral  counts,  apparently  have  intense  local  differentiation. 
The  variation  in  ventral  counts  has  been  discussed  in  detail  above 
(p.  177).  To  summarize,  twelve  females  from  Borneo  range  from 
137-200,  but  within  a  restricted  area  five  females  have  a  ventral 
range  of  11  (170-180).  Similarly,  five  males  from  the  same  locality 
have  counts  of  154-165,  whereas  twelve  males  from  scattered  parts 
of  Borneo  have  a  range  of  38  (144-181).  Our  Sumatran  sample 
has  the  same  pattern  of  variation. 

Variation  in  coloration  is  independent  of  that  in  ventral  counts 
and  does  not  show  the  same  amount  of  local  differentiation.  All 
fourteen  Bornean  adults  (over  300  mm.)  are  medium  to  light  brown 
above  with  the  outer  row  or  two  of  scales  slightly  lighter.  Two  out 
of  four  snakes  from  the  Malay  Peninsula  and  nine  out  of  ten  from 
Sumatra  are  dark  brown  or  almost  black  above  and  have  a  sharply 
defined  yellow  or  cream  stripe  on  the  two  outer  scale  rows.  The  ani- 
mals of  the  last  two  regions  therefore,  are  distinctly  bicolored, 
whereas  those  from  all  parts  of  Borneo  have  a  uniform  appearance. 

The  lined  coloration  of  grayi  forms  the  third  major  pattern  of 
vermiformis  and  it  is  characteristic  of  all  samples  from  the  southern 
Philippines.  This  lined  pattern  also  appears  in  the  highlands  of 
the  Malay  Peninsula  (Tweedie,  1950)  and  is  the  coloration  of  two 
of  the  Malayan  and  one  of  the  Sumatran  snakes  seen.  We  do  not 
have  sufficient  material  to  be  sure  that  whole  local  populations  of 
the  last  two  regions  are  characterized  by  this  pattern. 

Ventral  coloration  also  varies  geographically.  The  Bornean 
snakes  have  yellow  under  sides  crossed  by  black  bars  occupying  the 
complete  widths  of  2  to  4  ventrals;  occasionally,  a  few  of  the  cross- 
bars are  broken  up  into  alternating  black  squares.  The  same  ventral 
pattern  is  found  in  two  of  the  Malayan  and  three  of  the  Sumatran 


MARX  AND  INGER:  SNAKES  OF  GENUS  CALAMARIA         187 

snakes.  The  two  striped  Malayan  specimens  have  crossbars  that 
cover  4  to  6  ventrals.  The  other  seven  Sumatran  animals  have  the 
posterior  four-fifths  or  more  of  the  belly  solid  black  with  only  a  few 
narrow  crossbars  anteriorly. 

The  striped  snakes  from  the  Malay  Peninsula  and  Sumatra 
have  lower  ventral  counts  (two  males  154-157;  one  female  179) 
than  observed  in  grayi  (males  165-176;  females  192-207).  As  noted 
above,  grayi  differs  from  the  other  striped  vermiformis  in  the  number 
of  scales  in  contact  with  the  parietals.  The  similarity  of  grayi 
to  these  striped  individuals  is  probably  the  result  of  parallel  mutation 
in  similar  genetic  systems. 

On  the  basis  of  the  data  now  available  we  recognize  the  following 
forms  of  vermiformis: 

Calamaria  vermiformis  vermiformis  Dumeril  and  Bibron 

Calamaria  vermiformis  Dumeril  and  Bibron,  1854,  Erp.  Gen.,  7:  85 — Java. 
Calamaria  bruegeli  Mertens,  1924,  Zool.  Anz.,  60:  158 — central  Borneo. 

Descriptive  notes. — Borneo:  twelve  males;  total  length  160-436 
mm.;  tail  0.08-0.10  of  total;  ventrals  144-181  (mean  161.0±3.1; 
N=  12) ;  subcaudals  19-26  (mean  23.2±0.5;  N=  12) .  Twelve  females; 
total  length  120-642  mm.;  tail  0.06-0.08  of  total;  ventrals  137-200 
(mean  172.4±4.5;  N=12);  subcaudals  14-20  (mean  17.6±0.6; 
N=12). 

Sumatra:  four  males;  total  length  154-449  mm.;  tail  0.07-0.08 
of  total;  ventrals  166-196  (mean  186.8);  subcaudals  22-23  (mean 
22.8).  Seven  females;  total  length  186-503  mm.;  tail  0.05-0.06  of 
total;  ventrals  179-217  (mean  199.1±6.1;  N=7);  subcaudals  16-19 
(mean  17.3±0.4;  N=7). 

Malay  Peninsula:  two  males;  total  length  168-410  mm.;  tail 
0.10  (two)  of  total;  ventrals  154-157;  subcaudals  22-24.  Two 
females;  total  length  253-394  mm.;  tail  0.06  (two)  of  total;  ventrals 
179-183;  subcaudals  17  (two). 

Supralabials  5  or  6,  third  and  fourth  entering  the  eye;  infralabials 
5  or  6,  first  three  pairs  in  contact  with  anterior  chin  shields,  first 
pair  not  in  contact  behind  mental;  oculars  1+1;  frontal  V/2  to  23^ 
times  width  of  supraocular;  excluding  fifth  labial  each  parietal 
bordered  postero-laterally  by  3  (37)  or  4  (two)  scales;  position  of 
reduction  to  4  dorsal  scale  rows,  counting  subcaudals  from  terminal 
scute  0-9  (mean  4.3±0.3;  N=39);  maxilla  with  10  modified  teeth. 

Color  (in  alcohol)  varying  with  size;  small  specimens  (up  to  250 
mm.)  with  yellow  heads  and  narrow  yellow  rings  around  the  dark 


188  FIELDIANA:  ZOOLOGY,  VOLUME  37 

brown  body;  belly  yellow  barred  with  dark  brown;  large  specimens 
(over  340  mm.)  various  (see  above,  p.  186),  but  usually  brown 
above,  belly  with  black  crossbars. 

Discussion.— A  transition  between  these  two  color  patterns  is 
suggested  by  four  specimens  (234-375  mm.).  The  smallest  of  these 
has  the  yellow  body  rings  narrowed  and  interrupted  and  the  head 
somewhat  darkened  above.  The  largest  of  the  four  has  a  lighter 
head  than  usual  for  adults.  Gunther  (1865)  suggested  that  the 
yellow  rings  disappear  in  adults.  Smith  (1931)  also  comments  on 
the  difference  of  juvenile  and  adult  coloration. 

Earthworms  were  found  in  two  specimens. 

Localities  and  specimens  examined. — North  Borneo:  Jesselton 
District:  Mount  Kina  Balu,  Bundu  Tuhan,  4,500  feet  (USNM 
130252);  Mount  Kina  Balu,  Kiau,  3,000  feet  (BM  1929.12.22.110- 
111;  MCZ  43564-65;  RM  168-17);  Mount  Kina  Balu,  Kenokok, 
3,300  feet  (MCZ  43566-69).  Sandakan  District:  Sandakan  (CNHM 
15003). 

Sarawak:  First  Division:  Matang  (BM  1873.4.23).  Second 
Division:  Saribas  (SU  8575).  Fourth  Division:  Baram  District 
(BM  97.3.4-5,  1902.11.25.16-17);  Mount  Batu  Song,  1,000  feet 
(BM  92.10.7.8).  Fifth  Division:  Lawas  (CNHM  67278);  Pa 
Brayong  (CNHM  71646). 

Borneo:  Tanjong  (BM  96.2.17.11-12).  No  additional  data 
(BM  no  reg.  no.;  BM  72.2.19-55;  USNM  49802). 

Sumatra:  Kaba  Wetan  (USNM  70945-51);  Langkat  (MCZ 
37769-70);  Pandangsoke,  Bovenlanden  (MCZ  20971);  unknown  lo- 
cality (AMNH  2879). 

Malay  Peninsula:  Pahang:  Kuala  Terla,  4,500  feet  (RM 
4/1938);  Tanah  Rata,  4,600  feet  (RM  1930).  Selangor:  Bukit 
Lagong  (RM  8827);  Kepong  Forest  Reserve  (RM  13880). 

Calamaria  vermiformis  grayi  Gunther 

Calamaria  grayi  Gunther,  1858,  Cat.  Snakes  Brit.  Mus.,  p.  6 — Philippine 
Islands. 

Descriptive  notes. — Four  males;  total  length  149-358  mm.;  tail 
0.05-0.10  of  total;  ventrals  165-176  (mean  170.8);  subcaudals  22-27 
(mean  24.0) .  Four  females;  total  length  216-590  mm. ;  tail  0.04-0.06 
of  total;  ventrals  192-207  (mean  199.8);  subcaudals  15-18  (mean 
17.0). 


MARX  AND  INGER:  SNAKES  OF  GENUS  CALAMARIA         189 

Excluding  fifth  labial,  each  parietal  bordered  postero-laterally 
by  4  scales  in  all  but  one  animal;  position  of  reduction  to  4  dorsal 
scale  rows,  counting  subcaudals  from  terminal  scute  3-6  (mean 
5.0±0.4;  N=8);  other  counts  as  in  v.  vermiformis. 

Discussion. — Gunther  (1858)  considered  grayi  to  be  related  to 
lumbricoidea  Boie.  Reasons  are  given  above  for  relating  it  instead 
to  vermiformis. 

Localities  and  specimens  examined. — Philippine  Islands:  Basi- 
lan:  near  Isabela  (MCZ  43649).  Mindanao:  Cotabato  Province, 
Saub  (MCZ  25778-80;  USNM  120811);  Davao  Province,  Mount 
Apo,  2,800  feet  (CNHM  53373);  Lanao  Province,  Lumbatan 
(CNHM  15010).    Leyte:  Cabalian  (MCZ  25781). 

Calamaria  grabowskyi  Fischer 

Calamaria  grabowskyi  Fischer,  1885,  Arch.  Naturg.  Berlin,  51:  50,  pi.  4,  fig. 
1 — Telang  and  Tameanglaijang,  Dusson  Timor  District,  Borneo. 

Descriptive  notes. — Two  males;  total  length  320-340  mm.;  tail 
0.09-0.10  of  total;  ventrals  159-175;  subcaudals  26-27.  Two  fe- 
males; total  length  278-280  mm.;  tail  0.08-0.09  of  total;  ventrals 
152-183;  subcaudals  22-23. 

Supralabials  5,  third  and  fourth  entering  eye;  infralabials  5, 
first  pair  not  in  contact  behind  mental,  first  three  pairs  in  contact 
with  anterior  chin  shields;  oculars  1+1;  position  of  reduction  to  4 
dorsal  scale  rows,  counting  subcaudals  forward  from  terminal  scute 
14-19;  maxilla  with  9  modified  teeth. 

Color  (in  alcohol)  above  dark  brown  with  small  black  spots 
scattered  over  dorsal  scales;  center  of  scales  of  outer  row  light  yellow, 
lower  edge  of  first  scale  row  and  adjacent  edges  of  ventrals  dark 
brown;  ventrals  uniform  yellowish  in  anterior  eighth  of  body, 
followed  by  ventrals  with  black  anterior  edge  limited  at  first  to  a 
median  strip  then  widening  to  form  complete  dark  transverse  bars 
across  width  of  belly;  posterior  ventrals  uniform  yellow;  under  side 
of  tail  yellow  with  a  distinct  median  longitudinal  dark  stripe;  head 
above  dark  brown  with  black  spots;  upper  lip  yellowish;  head  below 
yellowish,  the  anterior  labials  edged  with  black. 

Discussion. — These  four  snakes  are  so  similar  to  descriptions 
(Fischer,  1885;  Boulenger,  1894)  of  grabowskyi  in  coloration,  in 
ventral  and  subcaudal  counts,  and  in  the  arrangement  of  the  head 
shields  that  we  have  no  doubts  about  their  identification.  It  should 
be  noted,  however,  that  a  number  of  named  forms  resemble  gra- 
bowskyi in  these  characters.    Among  these  forms  are  simalurensis, 


190  FIELDIANA:  ZOOLOGY,  VOLUME  37 

elegans,  prakkei,  and  suluensis,  all  of  which  differ  from  grabowskyi 
in  lacking  dark  ventral  bars. 

Localities  and  specimens  examined. — Sarawak:  Third  Division: 
Rejang  River,  Sarekei  (CNHM  67277).  Fourth  Division:  Bario 
(RM  no  numbers).  Fifth  Division:  La  was  District,  Pa  Brayong 
(CNHM  71600). 

Calamaria  suluensis  Taylor 

Calamaria  suluensis  Taylor,  1922,  Snakes  Phil.  Isls.,  p.  189 — Cagayan  Sulu. 

Descriptive  notes. — Two  females;  total  length  257-295  mm.;  tail 
0.09-0.11  of  total;  ventrals  148-168;  subcaudals  23-25;  supralabials 
5,  third  and  fourth  entering  eye;  infralabials  5,  first  pair  not  in 
contact  behind  mental,  first  three  pairs  in  contact  with  anterior 
chin  shields;  oculars  1+1;  position  of  reduction  to  4  dorsal  scale 
rows,  counting  subcaudals  forward  from  terminal  scute  9  (two); 
maxilla  with  7-8  modified  teeth. 

Discussion. — These  snakes  agree  remarkably  with  the  only 
known  specimen  of  suluensis,  a  female  having  154  ventrals  and  25 
subcaudals.  Like  our  specimens,  suluensis  is  brown  above  with 
scattered  dark  spots.  The  outer  row  of  scales  is  white.  A  dark 
brown  stripe  covers  the  lower  edge  of  the  outer  scale  row  and  the 
adjacent  edges  of  the  ventrals.  The  head  is  brown  with  dark  spots 
above,  the  upper  lip  is  yellow,  and  the  lower  lip  and  chin  are  yellow 
with  brown  spots.    The  belly  is  uniformly  yellow. 

As  indicated  in  the  discussion  of  grabowskyi,  suluensis  is  similar 
to  it,  and  to  simalurensis,  elegans,  and  prakkei.  The  ventral  counts 
of  suluensis  (females  148-168)  are  higher  than  those  of  simalurensis 
(136,  sex  unknown),  elegans  (133,  sex  unknown),  and  prakkei 
(126-144,  sex  unknown),  but  without  additional  data  for  sexed 
animals  these  differences  cannot  be  evaluated. 

In  addition  to  the  striking  distinction  in  ventral  coloration, 
suluensis  (as  we  understand  it)  differs  from  grabowskyi  in  the  fol- 
lowing characters: 

In  grabowskyi  the  postocular  is  elongate,  higher  than  wide,  its 
height  equal  to  its  distance  from  the  mouth,  and  as  high  as  the 
preocular.  In  suluensis  the  postocular  is  squarish,  its  height  about 
one-half  its  distance  from  the  mouth,  and  about  two-thirds  as  high 
as  the  preocular. 

The  parietals  of  grabowskyi  are  only  equal  in  length  to  the 
frontals,  whereas  in  suluensis  the  parietals  are  one  and  one-half 
times  the  length  of  the  frontal. 


MARX  AND  INGER:  SNAKES  OF  GENUS  CALAMARIA         191 

The  reduction  to  4  dorsal  scale  rows  takes  place  in  grabowskyi 
opposite  the  fourteenth  to  nineteenth  subcaudal  and  in  suluensis 
opposite  the  ninth  subcaudal,  in  both  cases  counting  the  subcaudals 
from  the  terminal  scute. 

Localities  and  specimens  examined. — North  Borneo:  Sandakan 
District:  Sandakan  (CNHM  63573).  Kudat  District:  Bongon  (BM 
1906.5.23.5). 

Calamaria  pendleburyi  Smith 

Calamaria  pendleburyi  Smith,  1931,  Bull.  Raffles  Mus.,  no.  5,  p.  27 — Kiau, 
Mount  Kina  Balu,  North  Borneo. 

Descriptive  notes. — One  male;  total  length  241  mm.;  tail  0.08  of 
total;  ventrals  136;  subcaudals  18.  Two  females;  total  length 
250-252  mm.;  tail  0.07  (one)  of  total;  ventrals  144-147;  subcaudals 
17  (one). 

Supralabials  5,  third  and  fourth  entering  eye;  infralabials  5, 
first  pair  not  in  contact  behind  mental,  first  three  pairs  in  contact 
with  anterior  chin  shields;  oculars  1+1;  position  of  reduction  to  4 
dorsal  scale  rows,  counting  subcaudals  forward  from  terminal  scute 
9-12;  maxilla  with  8  modified  teeth. 

Discussion. — Available  descriptions  of  margaritophora,  suma- 
trana,  and  curta  indicate  forms  similar  to  pendleburyi.  These  forms 
have  in  common  a  longitudinal  light  stripe  along  the  outer  row  of 
scales,  the  first  infralabials  not  in  contact  behind  the  mental,  the 
frontal  about  twice  as  wide  as  the  supraocular,  a  moderate-sized 
eye,  and  less  than  160  ventrals. 

Localities  and  specimens  examined. — North  Borneo:  Jesselton 
District:  Mount  Kina  Balu,  Bundu  Tuhan,  4,500  feet  (USNM 
130237) ;  Mount  Kina  Balu,  Kiau,  3,000  feet  (RM  146.17,  paratype); 
Mount  Kina  Balu,  Tenompok,  4,700  feet  (RM  18820). 

Calamaria  lautensis  de  Rooij 

Calamaria  lautensis  de  Rooij,  1917,  Rept.  Indo-Austr.  Arch.,  2:  163,  fig.  66 — 
Pulau  si  Laut,  Cocos  Island  and  Simalur. 

Descriptive  notes.— One  male;  total  length  142  mm.;  tail  0.09  of 
total;  ventrals  140;  subcaudals  19.  One  female;  total  length  224 
mm.;  tail  0.06  of  total;  ventrals  154;  subcaudals  12. 

Supralabials  5,  third  and  fourth  entering  eye;  infralabials  5, 
first  three  pairs  in  contact  with  anterior  pair  of  chin  shields,  first 
pair  of  infralabials  in  contact  behind  mental;  oculars  1+1;  position 


192  FIELDIANA:  ZOOLOGY,  VOLUME  37 

of  reduction  to  4  dorsal  scale  rows,  counting  subcaudals  forward 
from  terminal  scute  6-8;  maxilla  with  6-8  modified  teeth. 

Discussion.— Superficially  this  snake  resembles  many  described 
forms  (crassa,  suluensis,  tropica,  pendleburyi,  stahlknechti,  margarito- 
phora,  prakkei,  sumatrana,  elegans,  simalurensis,  muelleri,  bicolor, 
and  lautensis)  with  low  ventral  and  subcaudal  counts,  with  moderate- 
sized  eye,  and  with  frontal  about  twice  the  width  of  the  supraocular. 
In  coloration  these  forms  have  in  common  an  incomplete  neck 
band,  light  centers  to  the  outer  row  of  scales,  and  ventrals  with 
dark  outer  ends.  Only  bicolor  and  lautensis  have  the  first  infra- 
labials  in  contact  behind  the  mental.  Using  this  character  we  have 
tentatively  limited  identification  to  these  two  forms  and  have 
eliminated  bicolor  on  the  basis  of  the  size  of  the  eye  (smaller  in 
bicolor)  and  the  relative  width  of  the  frontal  (broader  in  bicolor) . 

The  gut  of  one  specimen  contained  an  earthworm. 

Localities  and  specimens  examined. — North  Borneo:  Jesselton 
District:  Mount  Kina  Balu,  Bundu  Tuhan,  4,500  feet  (USNM 
130236);  Mount  Kina  Balu,  Tenompok,  4,700  feet  (RM  295). 

Calamaria  everetti  Boulenger 

Calamaria  everetti  Boulenger,  1893,  Proc.  Zool.  Soc.  London,   1893:  525 — 
Sarawak. 

Descriptive  notes. — Two  females;  total  length  168-295  mm.  (latter 
with  incomplete  tail);  tail  0.11  (one)  of  total;  ventrals  151-171; 
subcaudals  20  (one);  supralabials  5,  third  and  fourth  entering  eye; 
infralabials  5,  first  infralabials  in  contact  with  each  other,  first  three 
infralabials  in  contact  with  anterior  chin  shields;  oculars  1+1; 
position  of  reduction  to  4  dorsal  scale  rows,  counting  subcaudals 
forward  from  terminal  scute  11;  maxilla  with  8-9  modified  teeth. 

These  specimens  agree  completely  with  the  original  description. 

The  Bornean  snake  was  found  on  the  ground  in  a  large  clearing. 

Localities  and  specimens  examined. — North  Borneo:  Kina- 
batangan  District:  southeast  end  of  Dewhurst  Bay  (CNHM  63572). 

Philippine  Islands:  Palawan:  Brooke's  Point,  Kabalnecan, 
1,400  feet  (CNHM  53381). 

Calamaria  brachyura  Boulenger 

Calamaria  brachyura  Boulenger,  1895,  Ann.  Mag.  Nat.  Hist.,  (6),  16:  481 — 
Mount  Kina  Balu,  North  Borneo. 

Descriptive  notes.— One  female;  total  length  258  mm.;  tail  0.03 
of  total;  ventrals  205;  subcaudals  10;  supralabials  5,  third  and 


MARX  AND  INGER:  SNAKES  OF  GENUS  CALAMARIA         193 

fourth  entering  eye;  infralabials  5,  first  pair  in  contact  behind  mental, 
first  three  pairs  in  contact  with  anterior  chin  shields;  oculars  1  +  1; 
position  of  reduction  to  4  dorsal  scale  rows,  counting  subcaudals 
forward  from  terminal  scute  3;  maxilla  with  8  modified  teeth. 

In  coloration  agreeing  with  the  original  description  except  for 
the  absence  of  yellow  spots  on  the  prefrontals. 

Found  under  a  log  in  a  cut-over  forest. 

Locality  and  specimen  examined. — North  Borneo:  Sandakan 
District:  Sapagaya  Forest  Reserve,  50  feet  (CNHM  63571). 

Calamaria  leucocephala  DumeYil  and  Bibron 

Calamaria  leucocephala  Dum6ril  and  Bibron,  1854,  Erp.  Gen.,  7:  83 — type 
locality  unknown. 

Descriptive  notes. — Two  males;  total  length  210-322  mm.;  tail 
0.16  (two)  of  total;  ventrals  143-150;  subcaudals  34-37.  Two 
females;  total  length  337-343  mm.;  tail  0.10  (two)  of  total;  ventrals 
161-169;  subcaudals  26-29. 

Supralabials  5,  third  and  fourth  entering  eye;  infralabials  5,  first 
pair  in  contact  behind  mental,  first  three  pairs  in  contact  with 
anterior  chin  shields;  oculars  1+1  and  0+1;  position  of  reduction 
to  4  dorsal  scale  rows,  counting  subcaudals  forward  from  tip  of  tail 
8-15;  maxilla  with  10  modified  teeth. 

Discussion. — These  specimens  agree  with  all  published  descrip- 
tions of  this  form.  The  preocular,  when  present,  is  very  small.  In 
one  specimen  it  is  lacking  on  both  sides,  and  in  a  second  is  present 
on  the  left  side  only.  Both  of  these  snakes  are  females.  In  all 
four  animals  the  prefrontals  enter  the  orbit. 

One  specimen  contained  the  remnants  of  an  earthworm. 

Locality  and  specimens  examined. — Locality  uncertain,  probably 
Singapore  Island  (CNHM  69976-79). 

Calamaria  pavimentata  DumeYil  and  Bibron 

Calamaria  pavimentata  Dumeril  and  Bibron,  1854,  Erp.  Gen.,  7:  71 — Java. 

Descriptive  notes. — One  male;  total  length  244  mm.;  tail  0.09  of 
total;  ventrals  160;  subcaudals  21;  supralabials  4,  second  and  third 
entering  eye;  infralabials  5,  first  pair  in  contact  behind  mental,  first 
three  pairs  in  contact  with  anterior  chin  shields;  oculars  1  +  1; 
position  of  reduction  to  4  dorsal  scale  rows,  counting  subcaudals 
forward  from  terminal  scute  5;  maxilla  with  8  modified  teeth. 


194  FIELDIANA:  ZOOLOGY,  VOLUME  37 

Locality  and  specimen  examined. — Indo-China:  Annam:  Bana 
(CNHM  11528). 

Calamaria  septentrionalis  Boulenger.    Figures  23,  25. 

Calamaria  septentrionalis  Boulenger,  1890,  Proc.  Zool.  Soc.   London,  1890: 
34 — Kiukiang  and  Hong  Kong. 

Descriptive  notes. — One  male;  total  length  168  mm.;  tail  0.07  of 
total;  ventrals  160;  subcaudals  18.  Two  females;  total  length 
277-342  mm.;  tail  0.03  (two)  of  total;  ventrals  179-180;  subcaudals 
10-11. 

Supralabials  4,  second  and  third  entering  eye;  infralabials  5, 
first  pair  in  contact  behind  mental,  first  three  pairs  in  contact  with 
anterior  chin  shields;  oculars  1+1;  dorsal  scale  rows  reduce  from  6 
to  1  abruptly  at  end  of  tail;  maxilla  with  8  modified  teeth. 

Two  specimens  had  eaten  earthworms. 

Locality  and  specimens  examined. — China:  Anhwei  Province: 
Ningkwo  (CNHM  7139-41). 

Calamaria  borneensis  Bleeker 

Calamaria  borneensis  Bleeker,  1860,  Nat.  Tijds.  Ned.  Indie,  21:  296 — Sintang, 
Borneo. 

Descriptive  notes. — Five  males;  total  length  135-279  mm.;  tail 
0.09-0.11  of  total;  ventrals  152-169  (mean  160.8);  subcaudals  22-24. 
Three  females;  total  length  184-360  mm.;  tail  0.06-0.08  of  total; 
ventrals  173-182;  subcaudals  16-19. 

Supralabials  4,  second  and  third  entering  eye;  infralabials  4, 
first  pair  not  in  contact  behind  mental,  first  two  pairs  in  contact 
with  anterior  chin  shields;  oculars  1+1;  position  of  reduction  to  4 
scale  rows,  counting  subcaudals  forward  from  terminal  scute  4-9; 
maxilla  with  8-9  modified  teeth. 

Color  (in  alcohol)  dark  grayish  brown  above  with  black  crossbars 
or  spots  mid-dorsally,  lateral  scale  rows  lighter;  belly  yellowish  with 
varying  amounts  of  black  in  the  form  of  checkers. 

Discussion. — A  single  specimen  from  Kuching  differs  from  the 
above  coloration  in  having  longitudinal  dark  lines  above  and  on 
the  lateral  edges  of  the  ventrals  and  in  lacking  the  dark  transverse 
markings.  Except  for  the  lateral  dark  lines  and  a  median  longi- 
tudinal dark  line  under  the  tail  this  snake  is  uniformly  yellow  below. 

Most  of  this  series  differ  from  descriptions  of  borneensis  (Bou- 
lenger, 1894;  de  Rooij,  1917)  in  the  presence  of  the  dark  transverse 


MARX  AND  INGER:  SNAKES  OF  GENUS  CALAMARIA         195 

markings  of  the  back.  However,  the  distinctive,  prominent  snout, 
the  checkered  belly,  and  the  counts  support  the  identification.  It 
is  clear  that  this  form  is  closely  related  to  linnaei  but  differs  from 
the  latter  in  the  shape  of  the  snout,  which  in  linnaei  is  rounded 
and  does  not  project  (Boulenger,  1894)  and  in  the  subcaudal  counts. 
Boulenger  (1894)  gives  the  subcaudal  counts  of  twelve  female 
linnaei  from  Java  as  9-15,  of  which  only  two  have  more  than  12. 

Javan  linnaei  show  the  same  variation  in  coloration  (Boulenger, 
1894)  as  is  indicated  for  borneensis  by  a  comparison  of  our  material 
with  the  description  given  by  Boulenger. 

Localities  and  specimens  examined. — Sarawak:  First  Division: 
Kuching  (RM,  no  number).  Fourth  Division:  Long  Mujan  (RM 
41,  44,  48-49,  73,  75;  SU  8569). 

Calamaria  buchi  sp.  nov.    Figure  26. 

Type. — Chicago  Natural  History  Museum  no.  71697,  a  female, 
from  Dalat,  Viet  Nam,  Indo-China.  Altitude  1,500  feet.  Collected 
by  Father  Buch  in  1953. 

Diagnosis. — A  Calamaria  with  the  eye  about  equal  to  its  distance 
from  the  mouth;  4  supralabials,  the  second  and  third  entering  the 
eye;  preocular  present;  frontal  about  twice  as  wide  as  supraocular; 
221  ventrals. 

Description. — Rostral  higher  than  broad,  visible  from  above; 
prefrontals  large,  maximum  length  subequal  to  length  of  frontal, 
posterior  border  angular,  in  contact  with  first  and  second  supra- 
labials and  entering  orbit  at  a  point;  preocular  present,  small, 
triangular;  postocular  small,  its  height  about  half  its  distance  from 
mouth;  supraocular  broad;  frontal  slightly  longer  than  broad,  less 
than  twice  width  of  supraocular,  about  two-thirds  length  of  parietals; 
4  supralabials,  second  and  third  entering  eye,  second  larger  than 
first  or  third,  fourth  largest;  first  infralabials  not  in  contact  with 
each  other;  mental  and  first  three  infralabials  in  contact  with 
anterior  chin  shields,  the  latter  in  contact  for  entire  length;  posterior 
chin  shields  about  two-thirds  length  of  anterior  ones,  in  contact 
anteriorly  only;  vertebral  row  of  nuchals  smaller  than  adjacent 
rows;  maxilla  with  9  modified  teeth. 

Scales  in  13  rows;  ventrals  221;  anal  entire;  subcaudals  13;  tail 
ending  in  a  blunt  point;  position  of  reduction  to  4  dorsal  scale  rows, 
counting  subcaudals  forward  from  terminal  scute  3. 

Color  (in  alcohol)  purplish  gray  above,  an  obscure  dark  vertebral 
line  anteriorly;  an  interrupted  light  nuchal  band;  outer  row  of 


196 


FIELDIANA:  ZOOLOGY,  VOLUME  37 


Fig.  26.    Calamaria  buchi  sp.  nov.  (type,  CNHM  71697);  actual  length  of 
head  7.5  mm. 


scales  white,  edged  above  with  black;  second  row  of  scales  white  in 
lower  half,  in  posterior  third  of  body  becoming  dark;  ventral  surface 
white,  uniform  except  for  dark  spots  on  infralabials  and  irregular 
dark  speckling  under  tail. 

Total  length  389  mm.;  tail  length  15  mm.,  length  of  head  to 
end  of  parietals  7.5  mm. 

Comparisons. — The  number  of  labials,  which  we  find  to  be  a 
relatively  unvarying  character,  and  the  high  ventral  count  eliminate 
all  but  javanica  lineata,  gimletti,  ventralis,  lowi,  and  gracillima  from 
consideration.  Calamaria  gracillima  has  more  than  300  ventrals,  no 
preocular,  and  a  tiny  supraocular;  the  first  lower  labials  are  in 
contact.  C.  lowi  has  198-236  ventrals  but  differs  from  buchi  in  the 
lack  of  preoculars,  in  the  smaller  eye,  in  the  relatively  broader 
frontal,  and  in  coloration;  the  entire  ventral  surface  of  lowi  is  dark; 
furthermore,  only  the  third  labial  enters  the  eye  in  lowi.  C.  ventralis 
has  249  ventrals,  but  has  no  preocular,  a  smaller  eye,  and  dark 
ventrals.     C.  gimletti  has  200-249  ventrals,  but  no  preocular,  a 


MARX  AND  INGER:  SNAKES  OF  GENUS  CALAMARIA         197 

broader  frontal,  the  first  lower  labials  in  contact,  and  the  outer 
rows  of  scales  dark.  C.  javanica  lineata  has  only  199  ventrals,  no 
preocular,  a  smaller  eye,  and  light  dorsal  stripes. 

Remarks. — The  gut  of  the  type  specimen  contained  the  remains 
of  an  earthworm. 

Calamaria  schmidti  sp.  nov.    Figure  27. 

Type. — United  States  National  Museum  no.  130240.  A  female 
from  Bundu  Tuhan,  Mount  Kina  Balu,  North  Borneo.  Collected 
at  4,500  feet,  on  August  1,  1951,  by  D.  H.  Johnson.  This  species  is 
named  in  honor  of  Karl  P.  Schmidt. 

Diagnosis.— A  Calamaria  with  the  eye  much  smaller  than  its 
distance  from  the  mouth;  4  supralabials,  the  first  three  subequal; 
no  preocular;  frontal  about  five  to  six  times  as  wide  as  supraocular; 
maxillary  teeth  conical;  first  pair  of  infralabials  in  contact  behind 
mental. 

Description  of  type. — Rostral  broader  than  high,  visible  from 
above;  prefrontals  squarish,  maximum  length  subequal  to  length  of 
frontal,  posterior  border  transverse,  in  contact  with  first  and  second 
labial  and  eye;  nasal  large,  slightly  larger  than  eye;  eye  small,  di- 
ameter one-half  its  distance  from  the  mouth;  no  preocular;  one  small 
postocular,  not  as  high  as  eye;  supraocular  small,  about  equal  to 
nasal;  frontal  about  as  wide  as  long,  pentagonal,  5  to  6  times  width 
of  supraocular,  about  two-thirds  length  of  parietals;  4  supralabials, 
first  three  subequal,  fourth  twice  length  of  second,  second  and  third 
entering  eye;  5  infralabials,  first  pair  in  contact  behind  mental, 
fourth  largest;  two  pairs  of  chin  shields,  both  pairs  in  contact, 
anterior  in  contact  with  three  infralabials,  anterior  pair  larger  than 
posterior  pair;  parietals  bordered  posteriorly  by  three  nuchals; 
vertebral  row  of  nuchals  distinctly  smaller  than  para-vertebral  rows. 
Maxilla  with  7  conical,  slightly  curved  teeth  widely  spaced  at  the 
base. 

Scales  in  13  rows;  ventrals  144;  subcaudals  14;  anal  entire;  tail 
ending  in  a  blunt  point;  position  of  reduction  to  4  dorsal  scale  rows, 
counting  subcaudals  forward  from  terminal  scute  6. 

Color  (in  alcohol)  purplish  gray  above,  uniform;  head  without 
markings;  supralabials  same  color  as  back;  anterior  infralabials  and 
first  pair  of  chin  shields  purplish  gray,  remainder  of  under  side  of 
head  yellowish;  ventral  surface  without  markings;  anterior  ventrals 
yellowish,  belly  becoming  increasingly  more  purple  posteriorly,  but 


198 


FIELDIANA:  ZOOLOGY,  VOLUME  37 


Fig.  27.    Calamaria  schmidti  sp.  nov.  (type,  USNM  130240);  actual  length 
of  head  9  mm. 


lighter  than  dorsal  color;  under  side  of  tail  darker  than  belly  but 
slightly  lighter  than  dorsal  surface. 

Total  length  253  mm.,  tail  length  19  mm.,  length  from  tip  of 
snout  to  posterior  border  of  parietals  9  mm. 

Paratype. — Stanford  University  no.  8568  from  Mount  Kina 
Balu,  North  Borneo.  This  animal  is  in  several  pieces  and  somewhat 
dried.  However,  the  head  scutellation  is  distinct  and  unmistakably 
identical  to  that  of  the  type. 

The  following  data  are  all  that  can  be  gleaned  from  this  snake; 
subcaudals  14;  supralabials  4,  second  and  third  entering  eye,  first 
three  subequal;  first  pair  of  infralabials  in  contact  behind  mental, 
first  three  pairs  in  contact  with  anterior  chin  shields;  frontal  5  to  6 
times  width  of  supraocular;  no  preocular;  postocular  present;  position 
of  reduction  to  4  dorsal  scale  rows,  counting  subcaudals  from 
terminal  scute  7;  maxilla  with  6,  widely  spaced,  conical  teeth. 


MARX  AND  INGER:  SNAKES  OF  GENUS  CALAMARIA         199 

Comparisons. — Only  three  species  of  Calamaria  agree  with 
schmidti  in  having  four  supralabials,  the  first  infralabials  in  contact 
behind  the  mental,  and  no  preocular.  The  three  are  gimletti,  gracil- 
lima,  and  doerianense.  The  first  two  differ  radically  from  schmidti 
in  ventral  counts,  gimletti  having  202-249  and  gracillima  having 
300-320  as  compared  to  144  in  the  type  of  schmidti.  The  frontal- 
supraocular  ratio  of  schmidti  (5-6)  is  about  twice  that  of  gimletti 
(3-3^). 

Calamaria  doerianense  has  the  second  and  fourth  supralabials 
subequal  in  length  whereas  in  schmidti  the  first  three  supralabials 
are  subequal  and  much  smaller  than  the  fourth.  The  frontal-supra- 
ocular  ratio  of  schmidti  (5-6)  is  twice  that  of  doerianense  (2-3) . 

Remarks. — The  type  specimen  contained  an  earthworm. 

Calamaria  lowi  Boulenger 

Calamaria  lovii  Boulenger,  1887,  Ann.  Mag.  Nat.  Hist.,  (5),  19:  169,  fig. — 
Rejang  River,  Sarawak. 

Descriptive  notes. — Two  males;  total  length  122-239  mm.;  tail 
0.07  (two)  of  total;  ventrals  198-202;  subcaudals  19-20.  Two 
females;  total  length  279-316  mm.;  tail  0.04-0.05  of  total;  ventrals 
223  (two);  subcaudals  14  (two). 

Supralabials  4,  third  only  entering  the  eye;  infralabials  4,  first 
pair  not  in  contact  behind  mental,  first  two  pairs  in  contact  with 
anterior  chin  shields;  no  preocular;  postocular  1;  position  of  reduction 
to  4  dorsal  scale  rows,  counting  subcaudals  forward  from  terminal 
scute  3-6;  maxilla  with  8-9  modified  teeth. 

In  all  respects  these  snakes  agree  with  the  original  description. 

Locality  and  specimens  examined. — SARAWAK:  Fourth  Division, 
Long  Mujan  (RM  51,  55,  no  number;  SU  8525). 

CHECK  LIST 

Calamaria  acutirostris  Boulenger 

Calamaria  acutirostris  Boulenger,  1896,  Ann.  Mag.  Nat.  Hist.,  (6),  17:  394 — 
Loka,  Mount  Bonthain,  Celebes. 

Calamaria  albiventer  Gray 

Changulia  albiventer  Gray,  1834,  111.  Ind.  Zool.,  2,  pi.  86,  figs.  6-9 — Penang. 

Calamaria  albopunctata  Barbour 

Calamaria  albopunctata  Barbour,  1908,  Bull.  Mus.  Comp.  Zool.,  51:  319 — 
East  Indies. 


200  FIELDIANA:  ZOOLOGY,  VOLUME  37 

Calamaria  alidae  Boulenger 

Calamaria  alidae  Boulenger,  1920,  Ann.  Mag.  Nat.  Hist.,  (9),  5:  282— Lebong 
Tandai  in  Benkulen,  Sumatra. 

Calamaria  apraeocularis  Smith 

Calamaria  apraeocularis  Smith,  1927,  Proc.  Zool.  Soc.  London,  1927:  224 — 
Djikoro,  Mount  Bonthain,  Celebes. 

Calamaria  beccari  Peters 

Calamaria  beccarii  Peters,  1872,  Ann.  Mus.  Civ.  Genova,  3:  34 — Sarawak. 

Calamaria  benjaminsi  Edeling 

Calamaria  benjaminsi  Edeling,   1864,  Ned.  Tijds.  Dierk.,  2:  202 — Marta- 
pura,  Borneo. 

Calamaria  bicolor  Dumeril  and  Bibron 

Calamaria  bicolor  Dumeril  and  Bibron,  1854,  Erp.  Gen.,  7:  78 — Borneo. 

Calamaria  bitorques  Peters 

Calamaria  bitorques  Peters,  1872,  Monatsber.  Akad.  Wiss.  Berlin,  1872:  585 
— Philippine  Islands. 

Calamaria  borneensis  Bleeker 

Calamaria  borneensis  Bleeker,  1860,  Nat.  Tijds.  Ned.  Indie,  21:  296 — Sintang, 
Borneo. 

Calamaria  brachyura  Boulenger 

Calamaria  brachyura  Boulenger,  1895,  Ann.  Mag.  Nat.  Hist.,  (6),  16:  481  — 
Mount  Kina  Balu,  North  Borneo. 

Calamaria  buchi  sp.  nov. 

Dalat,  Viet  Nam,  Indo-China. 

Calamaria  bungaroides  Werner 

Calamaria  bungaroides  Werner,   1901,   Zool.   Anz.,   24:   300 — type  locality 
unknown,  probably  Java. 

Calamaria  ceramensis  de  Rooij 

Calamaria  ceramensis  de  Rooij,  1913,  Bijdr.  Dierk.,  19:  26 — Honitetu,  Ceram. 


MARX  AND  INGER:  SNAKES  OF  GENUS  CALAMARIA         201 

Calamaria  collaris  Boulenger 

Calamaria  collaris  Boulenger,  1897,  Proc.  Zool.  Soc.  London,  1897:  225,  pi. 
14,  fig.  4 — Bone  Mountains,  Tomohon,  Rurukan,  Celebes. 

Calamaria  crassa  Lidth  de  Jeude 

Calamaria  crassa  Lidth  de  Jeude,   1922,   Zool.   Meded.,   6:  248 — Gunung 
Talamau,  Ophir  District,  Padang  Highlands,  Sumatra. 

Calamaria  curta  Boulenger 

Calamaria  curta  Boulenger,  1896,  Ann.  Mag.  Nat.  Hist.,  (6),  18:  62 — South 
Celebes. 

Calamaria  doderleini  Gough 

Calamaria  doderleini  Gough,  1902,  Zool.  Anz.,  25:  645 — Langkat,  Sumatra. 

Calamaria  doerianense  Brongersma 

Calamaria  doerianense  Brongersma,  1928,  Zool.  Anz.,  75:  255,  2  figs. — Pulau 
Durian,  Riouw  Archipelago. 

Calamaria  egregia  Barbour 

Calamaria  egregia  Barbour,  1927,  Proc.  Biol.  Soc.  Washington,  40:  127  — 
Mount  Tibang,  Borneo. 

Calamaria  electa  Barbour 

Calamaria  electa  Barbour,   1927,  Proc.  Biol.  Soc.  Washington,  40:   128 — 
Pasir,  Borneo. 

Calamaria  elegans  de  Rooij 

Calamaria  elegans  de  Rooij,  1917,  Rept.  Indo-Austr.  Arch.,  2:  158,  fig.  64 — 
Simalur  Island. 

Calamaria  everetti  Boulenger 

Calamaria  everetti  Boulenger,  1893,  Proc.  Zool.  Soc.  London,   1893:  525 — 
Sarawak. 

Calamaria  gervaisi  Dumeril  and  Bibron 

Calamaria  gervaisi  Dumeril  and  Bibron,  1854,  Erp.  Gen.,  7:  76 — Java  [in 
error,  see  text  p.  182]. 

Calamaria  gimletti  Boulenger 

Calamaria  gimletti  Boulenger,  1905,  Ann.  Mag.  Nat.  Hist.,  (6),  15:  456 — 
Kelantan,  Malay  Peninsula. 


202  FIELDIANA:  ZOOLOGY,  VOLUME  37 

Calamaria  goringi  Vogt 

Calamaria  goringi  Vogt,  1925,  Zool.  Anz.,  62:  64 — Java. 

Calamaria  grabowskyi  Fischer 

Calamaria  grabowskyi  Fischer,  1885,  Arch.  Naturg.  Berlin,  51:  50,  pi.  4,  fig. 
1 — Telang  and  Tameanglaijang,  Dusson  Timor  District,  Borneo. 

Calamaria  gracilis  Boulenger 

Calamaria  gracilis  Boulenger,  1896,  Ann.  Mag.  Nat.  Hist.,  (6),  18:  63 — 
Mount  Bonthain,  Celebes. 

Calamaria  gracillima  Gunther 

Calamaria  gracillima  Gunther,  1872,  Proc.  Zool.  Soc.  London,  1872:  594, 
pi.  39,  fig.  A — Sarawak. 

Calamaria  griswoldi  Loveridge 

Calamaria  lumbricoidea  griswoldi  Loveridge,  1938,  Proc.  Biol.  Soc.  Washington, 
51:  43 — Luidan  River,  near  Bundu  Tuhan,  Mount  Kina  Balu,  North 
Borneo. 

Calamaria  hollandi  Taylor 

Calamaria  hollandi  Taylor,  1923,  Phil.  Jour.  Sci.,  22:  550 — Port  Holland, 
Basilan,  Philippine  Islands. 

Calamaria  hosei  Gunther 

Calamaria  hosei  Gunther,  1896,  Ann.  Mag.  Nat.  Hist.,  (6),  17:  229 — Entoyut 
River,  Sarawak. 

Calamaria  indragirica  Schenkel 

Calamaria  indragirica  Schenkel,  1901,  Verh.  Ges.  Basel,  13:  164,  fig.  3 — 
Indragiri,  Sumatra. 

Calamaria  javanica  javanica  Boulenger 

Calamaria  javanica  Boulenger,  1891,  Ann.  Mag.  Nat.  Hist.,  (6),  7:  279 — 
Java. 

Calamaria  javanica  lineata  Brongersma 

Calamaria  javanica  var.  lineata  Brongersma,  1928,  Zool.  Anz.,  75:  256 — Long 
Petah,  Borneo. 

Calamaria  jeudei  Kopstein 

Calamaria  jeudei  Kopstein,  1926,  Zool.  Meded.,  9:  108 — Saparua  Island, 
Molucca  Islands. 


MARX  AND  INGER:  SNAKES  OF  GENUS  CALAMARIA         203 

Galamaria  joloensis  Taylor 

Calamaria  joloensis  Taylor,   1922,   Phil.  Jour.   Sci.,  21:  203 — Jolo   Island, 
Philippine  Islands. 

Calamaria  klossi  Smith 

Calamaria  klossi  Smith,  1926,  Ann.  Mag.  Nat.  Hist.,  (9),  18:  79 — Siberut 
Island,  Mentawai  Islands. 

Calamaria  lateralis  Mocquard 

Calamaria  lateralis  Mocquard,  1890,  Nouv.  Arch.  Mus.  Hist.  Nat.  Paris, 
(3),  2:  136,  pi.  8,  fig.  4— Mount  Kina  Balu,  Borneo. 

Calamaria  lautensis  de  Rooij 

Calamaria  lautensis  de  Rooij,  1917,  Rept.  Indo-Austr.  Arch.,  2:  163,  fig.  66 
— Pulau  si  Laut,  Cocos  Islands,  and  Simalur. 

Calamaria  leucocephala  Dumeril  and  Bibron 

Calamaria  leucocephala  Dumeril  and  Bibron,  1854,  Erp.  Gen.,  7:  83 — type 
locality  unknown. 

Calamaria  leucogaster  Bleeker 

Calamaria  leucogaster  Bleeker,  1860,  Nat.  Tijds.  Ned.  Indie,  21:  293 — Ampat- 
Lawang,  Sumatra. 

Calamaria  linnaei  Boie 

Calamaria  linnaei  Boie,  1827,  Isis,  20:  539 — Java. 

Calamaria  lowi  Boulenger 

Calamaria  lovii  Boulenger,  1887,  Ann.  Mag.  Nat.  Hist.,  (5),  19:  169,  fig. — 
Rejang  River,  Sarawak. 

Calamaria  lumbricoidea  Boie 

Calamaria  lumbricoidea  Boie,  1827,  Isis,  20:  540 — Java. 

Calamaria  lumholtzi  Andersson 

Calamaria  lumholtzii  Andersson,  1923,  Nyt.  Mag.  Naturk.  Kristiania,  61: 
123 — Tumbang  Maruwei,  Borneo. 

Calamaria  margaritophora  margaritophora  Bleeker 

Calamaria  margaritophora  Bleeker,  1860,  Nat.  Tijds.  Ned.  Indie,  21:  294 — 
Ampat-Lawang,  Sumatra. 


204  FIELDIANA:  ZOOLOGY,  VOLUME  37 

Calamaria  margaritophora  gastropicta  Holtzinger 

Calamaria  margaritophora  gastropicta  Holtzinger,  1920,  Arch.  Naturg.  Berlin, 
Abt.  85,  p.  106 — Indonesia. 

Calamaria  mearnsi  Stejneger 

Calamaria  mearnsi  Stejneger,  1907,  Smiths.  Misc.  Coll.,  50:  30 — Tangob, 
Mindanao,  Philippine  Islands. 

Calamaria  mecheli  Schenkel 

Calamaria  mecheli  Schenkel,  1901,  Verh.  Ges.  Basel,  13:  165,  fig.  4 — Indragiri, 
Sumatra. 

Calamaria  melanota  Jan 

Calamaria  linnaei  var.  melanota  Jan,  1862,  Arch.  Zool.  Anat.  Phys.,  2:  5 — 
type  locality  unknown. 

Calamaria  mindorensis  Boulenger 

Calamaria  mindorensis  Boulenger,  1895,  Ann.  Mag.  Nat.  Hist.,  (6),  16:  481 
— Mindoro,  Philippine  Islands. 

Calamaria  mjobergi  Lonnberg  and  Rendahl 

Calamaria  mjobergi  Lonnberg  and  Rendahl,   1925,  Ark.  Zool.  Stockholm, 
17A,  no.  23,  p.  1 — Brastagi,  Sumatra. 

Calamaria  muelleri  Boulenger 

Calamaria  muelleri  Boulenger,  1896,  Ann.  Mag.  Nat.  Hist.,  (6),  17:  394 — 
Loka,  Mount  Bonthain,  Celebes. 

Calamaria  nuchalis  Boulenger 

Calamaria  nuchalis  Boulenger,  1896,  Ann.  Mag.  Nat.  Hist.,  (6),   18:  62 — 
South  Celebes. 

Calamaria  occipitalis  Jan 

Calamaria  occipitalis  Jan,  1862,  Arch.  Zool.  Anat.  Phys.,  2:  9 — Java. 

Calamaria  ornata  Werner 

Calamaria  ornata  Werner,  1909,  Mitt.  Naturh.  Mus.  Hamburg,  26:  229, 
fig.  6 — Indragiri,  Sumatra. 

Calamaria  pavimentata  pavimentata  DumeYil  and  Bibron 

Calamaria  pavimentata  Dumeril  and  Bibron,  1854,  Erp.  Gen.,  7:  71 — Java. 


MARX  AND  INGER:  SNAKES  OF  GENUS  CALAMARIA         205 

Calamaria  pavimentata  formosana  Maki 

Calamaria  pavimentata  formosana  Maki,  1931,  Monog.  Snakes  Japan,  p.  136, 
pi.  51A,  figs.  90-91 — Arisan,  Formosa. 

Calamaria  pendleburyi  Smith 

Calamaria  pendleburyi  Smith,  1931,  Bull.  Raffles  Mus.,  no.  5,  p.  27 — Kiau, 
Mount  Kina  Balu,  North  Borneo. 

Calamaria  pfefferi  Stejneger 

Calamaria  pfefferi  Stejneger,  1901,  Proc.  Biol.  Soc.  Washington,  14:  191 — 
Miyako  Shima,  Yayeyama  group,  Riu  Kiu  Archipelago. 

Calamaria  polillensis  Taylor 

Calamaria  polillensis  Taylor,  1923,  Phil.  Jour.  Sci.,  22:  549 — Polillo,  Polillo 
Island,  Philippine  Islands. 

Calamaria  prakkei  Lidth  de  Jeude 

Calamaria  prakkei  Lidth  de  Jeude,  1893,  Notes  Leyden  Mus.,   15:  252 — 
Sandakan  Bay,  North  Borneo. 

Calamaria  quinquetaeniata  Despax 

Calamaria  quinquetaeniata  Despax,  1912,  Bull.  Mus.  Paris,   18:  203 — East 
Indies. 

Calamaria  raveni  Cochran 

Calamaria  raveni  Cochran,   1923,   Proc.  Biol.   Soc.  Washington,   36:  92 — 
Sungai  Merah,  Borneo. 

Calamaria  rebentischi  Bleeker 

Calamaria  rebentischi  Bleeker,  1860,  Nat.  Tijds.  Ned.  Indie,  21:  293 — Singa- 
wang,  Borneo. 

Calamaria  schlegeli  Dumeril  and  Bibron 

Calamaria  schlegeli  Dumeril  and  Bibron,  1854,  Erp.  Gen.,  7:  81 — Borneo. 

Calamaria  schmidti  sp.  nov. 

Bundu  Tuhan,  Mount  Kina  Balu,  North  Borneo. 

Calamaria  semiannulata  Boettger 

Calamaria  semiannulata  Boettger,  1898,  Kat.  Rept.  Senck.  Mus.,  2:  84 — 
Borneo. 


206  FIELDIANA:  ZOOLOGY,  VOLUME  37 

Calamaria  septentrionalis  Boulenger 

Calamaria  septentrionalis  Boulenger,  1890,  Proc.  Zool.  Soc.  London,  1890: 
34 — Kiu  Kiang  and  Hong  Kong. 

Calamaria  simalurensis  de  Rooij 

Calamaria  simalurensis  de  Rooij,   1917,  Rept.  Indo-Austr.  Arch.,  2:    159, 
fig.  65 — Simalur  Island. 

Calamaria  smithi  Dunn 

Calamaria  smithi  Dunn,  1923,  Jour.  Malayan  Asiat.  Soc,  1 :  3 — Long  Mujan, 
Sarawak. 

Calamaria  sondaica  Barbour 

Calamaria  sondaica  Barbour,  1908,  Bull.  Mus.  Comp.  Zool.,  51:  319 — Buiten- 
zorg,  Java. 

Calamaria  stahlknechti  Stoliczka 

Calamaria  stahlknechti  Stoliczka,  1873,  Jour.  Asiat.  Soc.  Bengal,  42:   119, 
pi.  11,  fig.  2— Dilli,  Sumatra. 

Calamaria  suluensis  Taylor 

Calamaria  suluensis  Taylor,  1922,  Snakes  Phil.  Isls.,  p.  189 — Cagayan  Sulu, 
Philippine  Islands. 

Calamaria  sumatrana  Edeling 

Calamaria  sumatrana   Edeling,   1870,   Nat.   Tijds.   Ned.   Indie,    31:   379 — 
Redjang,  Sumatra. 

Calamaria  tropica  Taylor 

Calamaria  tropica  Taylor,  1922,  Snakes  Phil.  Isls.,  p.  194 — near  Naujan, 
Mindoro,  Philippine  Islands. 

Calamaria  ulmeri  Sackett 

Calamaria  ulmeri  Sackett,  1940,  Not.  Nat.,  no.  41,  p.  2 — northwest  of  Blang- 
kedjeren,  Atjeh  Province,  Sumatra. 

Calamaria  uniformis  Smith 

Calamaria  pavimentata  var.  uniformis  Smith,  1921,  Proc.  Zool.  Soc.  London, 
1921 :  426 — Langbian  peaks,  Annam,  Indo-China. 

Calamaria  ventralis  Cochran 

Calamaria  ventralis  Cochran,  1923,  Proc.  Biol.  Soc.  Washington,  36:  91 — 
Sungai  Merah,  Borneo. 


MARX  AND  INGER:  SNAKES  OF  GENUS  CALAMARIA         207 

Calamaria  vermiformis  vermiformis  Dumeril  and  Bibron 

Calamaria  vermiformis  Dumeril  and  Bibron,  1854,  Erp.  Gen.,  7:  85 — Java. 

Calamaria  vermiformis  grayi  Gunther 

Calamaria  grayi  Gunther,  1858,  Cat.  Snakes  Brit.  Mus.,  p.  6 — Philippine 
Islands. 

Calamaria  virgulata  Boie 

Calamaria  virgulata  Boie,  1827,  Isis,  20:  540 — Java. 

Calamaria  zamboangensis  Leviton 

Calamaria  zamboangensis  Leviton,  1952,  Jour.  Washington  Acad.  Sci.,  42: 
239,  fig.  1 — Zamboanga,  Mindanao,  Philippine  Islands. 


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MARX  AND  INGER:  SNAKES  OF  GENUS  CALAMARIA         209 

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